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te mĀtauranga koiora biology BY3031 PLANT RESPONSES TO THEIR EXTERNAL ENVIRONMENT NCEA LEVEL 3 2013/1 biology ncea level 3 Expected time to complete work This work will take you about 20 hours to complete. You will work towards the following standard: Achievement Standard 91603 (Version 1) Biology 3.3 Demonstrate understanding of the responses of plants and animals to their external environment Level 3, External 5 credits In this booklet you will focus on these learning outcomes: •• describing the environment, adaptations and ecological niche •• discussing plant responses to the biotic environment •• discussing plant responses to the abiotic environment. You will continue to work towards this standard in booklet BY3032: Animal responses to their external environment. Copyright © 2013 Board of Trustees of Te Aho o Te Kura Pounamu, Private Bag 39992, Wellington Mail Centre, Lower Hutt 5045, New Zealand. All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any means without the written permission of Te Aho o Te Kura Pounamu. © te ah o o te k u ra p ou n am u contents 1 The environment: an overview 2 Competition and cooperation 3 Defence and cooperation 4 Orientation responses 5 Control of plant growth 6 Timing responses 7 Teacher-marked assignment 8 Answer guide Glossary © te ah o o t e k ur a p o un a m u BY3031 1 how to do the work When you see: Use the Topic webpage or the Internet. 1A Complete the activity. Check your answers. Contact your teacher. You will need: •• a pen and paper •• access to the Internet •• the teacher-marked assignment (BY3031A). Resource overview Throughout this booklet you are directed to many links to websites and videos. These provide useful and interesting information that will help to explain the subject content. You should aim to access these links. The assumption is that you allocate at least three hours to each lesson: one to two hours to view and work through these websites and one hour to work through each lesson. If you have not studied NCEA Level 2 Biology, contact your teacher to discuss this. In this booklet, you will read about a range of responses plants exhibit towards their biotic and abiotic environment. This booklet is for your own use and does not need to be sent in for marking. However, you are asked to send in your answers to the teacher-marked assignment (BY3031A) as well as your selfassessment. 2 BY3031 © te ah o o te k u ra p ou n a mu the environment: an overview learning intentions In this lesson you will learn to: •• describe the biotic and abiotic environment •• explain adaptations •• explain the ecological niche of an organism. introduction You have probably seen or experienced most of the following situations: •• the leaves of deciduous trees changing colour and falling during autumn •• insects drawn towards lights at night •• dogs growing thicker coats as winter approaches •• plants on the forest floor growing upwards towards the light •• a cat raising the hairs on its back and arching its back to appear larger when approaching a rival cat •• plants producing flowers in summer •• a snail retreating into its shell if threatened •• shellfish such as pipi and clams burrowing down into the sand when disturbed •• blue penicillin mould growing on old bread or fruit •• your stomach feeling as if it has ‘butterflies’ inside when you are nervous or excited. These are just a few of many examples of plants, animals and microorganisms responding to changes in their environments. Every species needs to be responsive to changes and stimuli in their surrounding environments and make suitable responses to these stimuli, in order to survive. A stimulus (plural: stimuli) is a change in the environment that causes a response in an organism. istockphoto.com 1 Fig 1.1: When people are excited or happy they are responding to their environment. Animals, being mobile, are able to respond quickly, usually by moving towards or away from a stimulus, to find more favourable conditions. Plants are usually not mobile, so they respond to stimuli by adjusting their growth and development. In this topic you will learn how plants and animals respond to environmental changes and stimuli, and how these responses optimise the plants’ and animals’ survival in their particular ecological niches. There are a large number of biological terms associated with this topic so we suggest you make up a glossary as you go through the work. This will help you learn and remember the many terms and their meanings. The glossary at the back of this booklet could provide you with a good start. © te ah o o t e k ur a p o un a m u BY3031 3 the environment: an overview the environment: an overview The environment surrounding a plant or animal in its habitat (the place where it lives) is made up of all the living organisms (the biotic environment), the non-living factors (the abiotic environment) and the nutrient cycles and energy flows that connect all of these. Together they form an ecosystem. biotic environmental factors These are the influences of living organisms on each other in a community. All the organisms of every species in any community interact with each other and influence each other. Some species provide shelter and protect others, while other species harm their neighbours in the community. The living organisms that make up the biotic environment act as stimuli to which other organisms in the environment need to respond. Biotic factors may be: •• intraspecific – between members of the same species, (‘intra’ means within) or •• interspecific – between members of different species, (‘inter’ means between). Some examples of intraspecific biotic factors are competition for resources such as: food, water, light, space, nesting sites and mates cooperation strategies within species for defence, hunting and survival aggressive interactions to establish hierarchies and defend territories Intraspecific competition can be intense, as all the individuals have the same requirements for resources. Interspecific competition occurs if different species have similar requirements for any resource. Some examples of interspecific biotic factors are competition for resources such as: food, water, light and space mutualism, which is an association between species for their mutual benefit exploitation of one species by another, such as parasitism and predator/prey interactions abiotic environmental factors These are physical factors that can act as stimuli to which the organisms need to respond. Factors like these are all non-living (abiotic) influences: soil type intensity and direction of light salinity and clarity of water availability and amount of water availability and quality of air pressure (altitude) the average and range of temperature velocity and direction of wind 4 BY3031 the effect of gravity the presence of chemicals and nutrients © te ah o o te k u ra p ou n a mu the environment: an overview adaptations In order to survive in its habitat and environment, every organism has special adaptations. An adaptation is any inherited characteristic that enables an organism to survive and reproduce in its habitat. Adaptations may be structural, behavioural or physiological. istockphoto.com Structural adaptations (Fig 1.2) are features of body structure. Some examples of structural adaptations are a snail’s shell for protection; a bird’s wing for flight; a cat’s claw to catch prey; a plant’s leaves to trap sunlight for photosynthesis; and a plant’s flower to attract pollinators. Fig 1.2: A cat’s claws are a structural adaptation to catch its prey. istockphoto.com Behavioural adaptations (Fig 1.3) are types of behaviour that help an organism to survive. Some examples of behavioural adaptations are a male bird feeding the female bird sitting on the nest; a hedgehog curling into a ball when threatened; a possum’s nocturnal activities; and slaters (woodlice) hiding under logs to escape from high temperatures on the soil surface during the day. Fig 1.3: Meerkats work as a team and stand guard. This behaviour protects the family group from predators. istockphoto.com Physiological adaptations (Fig 1.4) are often chemical or physical adaptations. For example, humans keep their body temperature the same (about 37ºC) despite changes in the temperature of the external environment; pale skin tans when exposed to sun over long periods; and callouses form on hands and feet in response to repeated contact or pressure. Fig 1.4: A snake producing venom to defend itself is a physiological adaptation. 1A adaptations and behaviours Go to the Topic webpage to learn more about adaptations and behaviours. © te ah o o t e k ur a p o un a m u BY3031 5 the environment: an overview ecological niche An organism’s ‘way of life’ or ecological niche encompasses the ways in which an organism carries out all its life processes. This includes its interactions with its biotic and abiotic environment, such as: •• its relationships with other organisms •• its reproductive strategies to ensure the species survives •• the resources and opportunities provided by the habitat •• and the adaptations it has which enable it to take advantage of these opportunities. The work of a Russian microbiologist and ecologist, Georgii Gause, led him to publish the principle of competitive exclusion. This principle became known as Gause’s principle. It states that no two species with identical ecological niches can co-exist for long in the same place. If there is fierce competition between species, one species may be eliminated. If the competition is only moderate the two species can co-exist in the same habitat because their ecological niches are slightly different. 1B ecological niches Go to the Topic webpage to learn more about ecological niches. tolerance Although plants and animals are adapted to their particular environments and ecological niches, they need to be able to adapt to slow changes in their environments or they may not survive otherwise. An organism’s ability to survive variation in environmental conditions is called its tolerance. Tolerance to change in environmental factors varies between individuals of the same species. Acclimatisation is the ability of an organism to adjust its tolerance limits or optimum range in response to slow changes in its environment. For example, mountain climbers who want to conquer high-altitude peaks normally live at low altitudes. They have to spend several days at middle altitudes to prepare their bodies for the decreased amount of oxygen at high altitudes. During this period of acclimatisation, their bodies increase the production of red blood cells (the cells that carry oxygen around the body). This is a physiological adaptation. This is just one example that shows how organisms can adjust their range of tolerance, provided the changes occur slowly. Most organisms can adapt to keep within their ‘optimum range’ of tolerance. If any factor in an environment is outside an organism’s range of tolerance, the organism cannot live in that environment. Constant monitoring of the environment to detect the stimuli that indicate changing environmental conditions is vital if an organism is to survive. 6 BY3031 © te ah o o te k u ra p ou n a mu the environment: an overview Look at Fig 1.5 below showing how the effect of different physical factors must be kept within a narrow optimum range. This range is known as the level of tolerance for each species. Lower limit of tolerance Optimum range Zone of physiological stress Zone of physiological stress Upper limit of tolerance preferred niche too cold too wet too acidic temperature -5°C 15°C 25°C 50°C moisture (%) of soil 5 10 3.0 4.0 20 30 40 50 pH 5.5 7.5 9 too hot too dry too alkaline 10 Fig 1.5: Level of tolerance of a daisy to three abiotic factors. Source of data: NZQA As you can see from the graph, the preferred niche of the plant has a narrow temperature, moisture and pH range. As conditions change, the plant needs to be able to detect the stimuli indicating change and respond to the changes in order to survive (zone of physiological stress). If the conditions become too extreme (beyond its lower or upper limit of tolerance), the plant will not survive. Plants and animals continually have to interact and respond to their environments in order to survive. The term ‘fitness’ is used to describe how well suited an organism is to survive in its habitat. © te ah o o t e k ur a p o un a m u BY3031 7 the environment: an overview 1C defining terms Define the following terms. Stimulus: Habitat: Biotic environment: Abiotic environment: Ecosystem: Intraspecific: Interspecific: 8 BY3031 © te ah o o te k u ra p ou n a mu the environment: an overview Adaptation: Tolerance: Acclimatisation: Ecological niche: Check your answers. the new zealand falcon The New Zealand falcon (Fig 1.6) is a bird of prey. New Zealand falcons hunt for live prey, mainly by watching from a vantage point and making a fast direct flying attack, flying at speeds of over 100 km/h and either striking or grasping the prey with their sharp talons. They kill their prey with a quick, powerful bite to the neck. They prey on insects, mammals and lizards, but their diet consists mainly of birds. rob suisted 1D Fig 1.6: New Zealand falcon/kārearea (Falco novaeseelandiae). The falcon has many adaptations to catch its prey while in flight (‘on the wing’). Explain at least two adaptations that the falcon has that suit it to its ecological niche. © te ah o o t e k ur a p o un a m u BY3031 9 the environment: an overview Adaptation 1: Adaptation 2: Explanation: Check your answers. key words key points stimulus habitat adaptations biotic environment abiotic environment ecological niche •• A stimulus is a change in the environment that causes a response in an organism. •• A habitat is the place where an organism lives. •• An adaptation is an inherited characteristic that enables an organism to survive and reproduce in its habitat. •• Adaptations may be structural, behavioural or physiological. •• The biotic environment is the influences of living organisms on each other. •• The abiotic environment is the non-living influences on an organism. •• The ecological niche is a description of how the organism feeds, the opportunities provided by the habitat and the adaptive features of the organism which enable it to take advantage of these opportunities. 10 BY3031 © te ah o o te k u ra p ou n a mu competition and cooperation learning intentions In this lesson you will learn to: •• discuss competitive plant–plant responses to the biotic environment •• discuss cooperative plant–plant responses to the biotic environment. introduction You may think that plants are not very exciting organisms – they grow, some flower, and some have leaves that change colour and fall in autumn, but you may not think of them as very responsive organisms. This is just not true! Plants are in fact extremely evolved organisms, acutely attuned to their environment. They are able to detect very subtle changes or stimuli in the environment and respond to these, in some cases with great rapidity. In this lesson you will learn how plants compete but also cooperate with each other, and what strategies they have evolved to get a competitive edge. competition for resources In any habitat, the plants all have basic requirements for resources such as light, water, space to grow, minerals from the soil and carbon dioxide from the atmosphere. If any of these resources are in short supply, the plants’ growth will be slowed. If many plants are growing close together in the same habitat and they have similar, or the same, resource requirements, there will be fierce competition among the plants for these resources. Whenever two niches overlap there will be competition between the organisms concerned. Those plants that are best suited to monopolise the resources will have a competitive advantage and will be more successful in growing and reproducing than their neighbours. In plants the competition is usually indirect, through the resource, rather than directly with the other plants. Competition may be intraspecific or interspecific and may be either through interference in the growth of other species or through exploitation of other species. sunlight Plants compete fiercely for sunlight, an essential requirement for photosynthesis. In a forest (Fig 2.1), taller plants receive more light and may block most of the light from reaching the forest floor. The plants on the forest floor have adapted to survive in lower light intensities, some growing larger leaves to compensate for the lack of light. Plants in a forest have adapted to different light intensities. The plant growth forms vertical layers, with those plants that demand a lot of light growing in the canopy and those Fig 2.1: A forest showing stratification. that require less light growing in the layers below. This vertical layering of plants is called stratification and is shown in the image on the right. © te ah o o t e k ur a p o un a m u BY3031 don laing 2 11 competition and cooperation The main way plants compete for sunlight is to grow towards the light, overshadowing other competing plants nearby, and slowing their growth. They are utilising the available resources more quickly than neighbouring plants. Other ways plants compete by better utilising the available resources are by growing longer roots or a more extensive root system to take advantage of the water supply, or being better able to tolerate adverse conditions. Note that while the available resources are abiotic environmental factors, the competition between plants for these resources is biotic, as the plants are living elements of the environment. seeds Other plants depend on birds or insects to Fig 2.2: A gorse plant on a Wellington hillside. spread their seeds so that the seedlings are able to grow in more favourable, less competitive conditions away from the parent plant. don laing When walking in the hills you may have heard the explosive ‘popping’ sounds that gorse seedpods (Fig 2.2) make as they break open and scatter their seeds over a large area. This is a strategy that the gorse plants have evolved to ensure their seeds are spread over a wide area and the seedlings will not compete with the parent plant or with each other. chemical competition Some plants produce chemicals that are toxic to other plants, so that the growth of nearby competing plants is inhibited. The production of toxic chemicals that inhibit growth is called allelopathy. The toxic chemicals leach from the roots into the surrounding soil or accumulate in the soil around the plant as the leaves drop and decay. Desert plants also use allelopathy as a means of ensuring no plants grow nearby so that their roots can use the scarce water resource to their advantage. Many other plants including wetland species, grasses, tobacco, rice and pea plants are known to produce root allelotoxins to reduce competition with neighbouring plants. 2A istockphoto.com An example is the walnut tree (Fig 2.3), which produces a chemical called juglone in its leaves and roots that inhibits the growth of other plants nearby. Fig 2.3: A walnut tree. allelopathy Go to the Topic webpage to learn more about allelopathy. 12 BY3031 © te ah o o te k u ra p ou n a mu competition and cooperation exploitation Some species actively exploit their neighbours in order to compete for resources. In the forest you may have noticed the long vines or lianas that grow up trees to reach the sunlight. These lianas are making use of (exploiting) the trunks of neighbouring trees to help them reach the sunlight they need for photosynthesis. Epiphytes such as some orchids perch high up on the branches of tall trees so that they can reach light of a higher intensity than on the forest floor and are protected from herbivores. rob suisted The beautiful native Northern rātā (Metrosideros robusta) (Fig 2.4) can start life as an epiphyte or seedling plant perched high on a host tree, usually a rimu. The rātā grows roots down to the ground, finally enclosing and strangling its host as it grows into a big tree. In this way the rātā exploits the host tree to establish itself and gain access to sunlight higher up in the forest canopy. Fig 2.4: The Northern rātā. rob suisted Mistletoe (Fig 2.5) is another example of a plant that exploits another by parasitising it. The mistletoe is in fact semi-parasitic, as it has green leaves that can photosynthesise but it relies on tapping into the xylem tissues of a host tree for water and nutrients. Fig 2.5: A mistletoe plant. 2B competition between plants Go to the Topic webpage to learn more about plant competition. © te ah o o t e k ur a p o un a m u BY3031 13 competition and cooperation plant cooperation Although competition between plants can be fierce, plants also cooperate with each other in order to survive. cooperative strategies Some plants find it beneficial to grow close together. In this way they provide support for each other to grow tall and reach the light. Wind-pollinated plants also use this strategy to reach the higher wind levels so that their pollen can be blown easily to other plants of the same species. 2C cooperative strategies Fig 2.6: Feijoa trees need another feijoa tree for pollination. istockphoto.com The roots of marigold plants (Tagetes species) (Fig 2.7) secrete a chemical that kills plant parasitic nematodes, a kind of roundworm that damages plants. The nematodes cause swellings on the roots, which stunt the plants’ growth and can lead to their death. Producing a toxic chemical is a form of allelopathy, but unlike in the previous examples, where allelopathic chemicals were detrimental to the growth of other plants, the chemical produced by marigolds is beneficial, as it kills the nematodes which can damage the plants. This is an example of allelopathic cooperation between plants. istockphoto.com Some fruit trees such as feijoas (Feijoa sellowiana) (Fig 2.6) cannot be self-pollinated, so they need another tree of the same species growing nearby for pollination. By growing close together, both trees benefit from crosspollination and can produce flowers and fruit with increased genetic diversity as a result. Fig 2.7: Marigold plants. Go to the Topic webpage to learn more about the association between marigolds and nematodes. 14 BY3031 © te ah o o te k u ra p ou n a mu Some species of plants actively benefit others growing nearby. For example, plants of the legume family (including peas, beans, clover, peanuts, alfalfa and soy) have rhizobium bacteria living in nodules on their roots (Fig 2.8). The rhizobium bacteria can convert nitrogen gas from the air into ammonium in the soil. This process is called nitrogen fixation because the rhizobium bacteria ‘fix’ the nitrogen by converting it from the unavailable form found in the atmosphere to a form which plants can use. The legume plants receive nitrogen in a useful form for growth and the rhizobium receives carbohydrates from the plant, produced by photosynthesis. The relationship between legumes and rhizobium bacteria is symbiotic or mutualistic (both organisms benefit). Although most of the useful nitrogen is removed when the legumes are harvested, significant amounts can remain in the soil for the use of future crops. te kura competition and cooperation Fig 2.8: Plant roots showing nodules on the roots where rhizobium bacteria live. plant–fungus relationships lichens te kura You have probably seen lichen (Fig 2.9) growing on trees or old fence posts. Lichen is actually not a plant but a symbiotic relationship between a fungus and an alga. The fungus’s hyphae (fine threads or filaments) provide a sheltered place for the alga as well as providing it with water and nutrients. In return the alga, which contains chlorophyll, carries out photosynthesis and provides carbohydrates for the fungus. Fig 2.9: Lichens growing on a tree. 2D lichens Go to the Topic webpage to learn more about lichens. did you know? Some lichens are used to create colours to dye for! New Zealand spinners who dye their own wool prize the lichen Pseudocyphellaria coronata. Compared with brightly coloured synthetic dyes, lichen dyes, some of which are pleasantly fragrant, yield a wide range of subtle colours depending on what mordant is used. Source: http://www.teara.govt.nz/en/lichens/2 © te ah o o t e k ur a p o un a m u BY3031 15 competition and cooperation mycorrhizal fungi Mycorrhizal fungi have a close and beneficial association with the roots of plants. Most plants co-exist with these fungi, which help them absorb nutrients from the surrounding soil. The fungal hyphae extend from the host plant into the surrounding soil. This increases the surface area for absorption of water and nutrients by both the fungus and the host plant. The fungi also secrete growth factors that stimulate the roots to grow and branch and may produce antibiotics that may protect the plant from pathogens. In return the fungus receives carbohydrates made by photosynthesis in the plant’s leaves. Mycorrhizal fungi are an important part of pine forest ecosystems. The mushrooms that appear above ground around the trees are the fruiting bodies of these fungi. 2E mycorrhizal fungi Go to the Topic webpage to learn more about mycorrhizal fungi. 2F competitive strategies 1. Define allelopathy and explain why this process gives plants a competitive advantage. Allelopathy: Explanation: 16 BY3031 © te ah o o te k u ra p ou n a mu competition and cooperation phil bendle 2. The perching lily, Collospermum hastatum, (kahakaha or ‘widow maker’) (Fig 2.10) is one of the largest epiphytes in the New Zealand bush, growing high in the branches of canopy trees. Perching lilies are tall, clump-forming plants with long, arching leaves. What competitive advantages do epiphytes such as the perching lily have compared to lower growing forest plants? Fig 2.10: Perching lilies. Check your answers. © te ah o o t e k ur a p o un a m u BY3031 17 competition and cooperation 2G plant cooperation Truffles, such as the black truffle, Tuber melanosporum, are an expensive fungal delicacy. These mycorrhizal fungi are now cultivated in New Zealand, on the roots of oak and hazel trees. Discuss why trees that have an association with mycorrhizal fungi tend to grow better than those which don’t have this association. Check your answers. key words key points competition exploitation allelopathy resources cooperation symbiotic association mutualistic association •• Competition may be intraspecific or interspecific and may be either through interference in the growth of other species or through exploitation of other species. •• Resources for which plants compete include sunlight, water, nutrients in the soil, carbon dioxide from the atmosphere and space to grow. •• Plants compete with each other for resources in order to survive in the environment. •• Plants have developed cooperative strategies that help them to survive. These may be symbiotic associations with other plants. 18 BY3031 © te ah o o te k u ra p ou n a mu defence and cooperation learning intentions In this lesson you will learn to: •• discuss defensive plant–animal responses to the biotic environment. •• discuss cooperative plant–animal responses to the biotic environment. introduction Plants are at the start of all food chains. They are eaten by herbivores, which are in turn eaten by carnivores. There are many kinds of herbivores: grazers such as sheep and cattle that eat pasture plants; browsers such as giraffes and elephants that eat leaves of shrubs and trees; fruit and seed eaters such as humans, bats and birds; sap suckers such as aphids; and nectar feeders such as birds and butterflies. However, plants generally prefer not to be eaten! As they cannot move away from their predators, they have developed strategies to avoid predation by many herbivores. physical defences If you have ever picked blackberries or roses from the garden you will know that some plants have thorns or spines. These are a very effective defence against herbivores that are likely to browse on the plant. The New Zealand native plant matagouri is an example of a plant with thorns. Many plants produce protective barriers such as bark or gum or thick waxy cuticles to prevent predation by herbivores. Others have evolved defensive growth habits. An example is divarication, where plants branch repeatedly and produce a tangle of stems with the outer branches having smaller and fewer leaves than the inner branches. The stems of divaricating plants are often tough and difficult to break. This adaptation makes it difficult for browsers to reach the inner, larger leaves. New Zealand has many divaricating plants, such as the Coprosma species (Fig 3.1). It is thought that the many divaricating species in New Zealand arose as an adaptation to browsing by moa. wikimedia commons 3 Fig 3.1: A Coprosma plant showing divarication. Some trees such as beech trees prevent their seeds being eaten by a process called seed masting. Every few years all the trees of the same species release all their seed simultaneously. This has two advantages for the trees: first, with so many seeds available as food, they won’t all be eaten by seed-eating animals and so some will survive to germinate. Secondly, as there are not enough seeds in the intervening years to support a large population of seed-eating animals, there is less chance of the seeds being eaten during a mast year. © te ah o o t e k ur a p o un a m u BY3031 19 defence and cooperation chemical defences Some plants prevent attack by herbivores by producing chemicals that are unpalatable, toxic or insect-resistant. Familiar herbs and spices such as lavender, thyme, rosemary, mint, cinnamon and cloves, for example, produce oils in their leaves (essential oils) that we enjoy in cosmetics and food, the real purpose of which is to discourage insect predators from eating their leaves. Substances such as morphine in opium poppies and nicotine in tobacco are other examples of toxic chemicals produced as plant protection. In New Zealand the toxin 1080 is commonly used against browsing possums that are destroying native forests. This is a human-made chemical reproduction of a naturally occurring, biodegradable toxin, sodium monofluoroacetate, found in many Australian, South American and South African plants such as eucalyptus. Low concentrations are also found naturally in tea (Camellia species). Plants have developed monofluoroacetate as a natural defence against browsing mammals in those countries. Some plants produce an unusual amino acid called canavanine, which is similar to the amino acid arginine, which insects need as part of their diet. When insects eat these plants, they incorporate canavanine instead of arginine into their proteins. This makes these proteins ineffective, and eventually the insects die. When willow trees are attacked by pathogens, they produce anti-microbial compounds called phytoalexins, which destroy the pathogens. The willows also produce other chemicals such as salicylic acid, which travel through the tree, triggering an anti-microbial response to the pathogens in leaves in other parts of the tree. Some plants go even further and release chemicals that attract predatory animals to prey on their attackers. These plants, when eaten by caterpillars, release volatile (airborne) chemicals that attract dragonflies that feed on the caterpillars. Others attract parasitic wasps that lay their eggs inside the caterpillars that are eating the plants’ leaves. The eggs hatch inside the caterpillars and eat their way out, killing the caterpillars in the process. The plants benefit from the destruction of the caterpillars. Some plants and animals produce life-saving chemicals. Certain chrysanthemums, for example, produce chemicals called pyrethrins in their leaves, which are very poisonous to insects and stop them from eating the leaves. However, the caterpillars of certain moths and butterflies have developed an enzyme that can detoxify (break down) the pyrethrins, and these caterpillars can eat the leaves unharmed (Fig 3.2 on next page). Some chrysanthemum species have gone one better, and produced another chemical called sesamin that can knock out the protective enzyme in these caterpillars. There certainly is chemical warfare in nature! 20 BY3031 © te ah o o te k u ra p ou n a mu defence and cooperation Pyrethrins (in powder form and as sprays) are used by gardeners for protecting plants against insects. Prevents most insects from attacking the plant. Detoxifying enzymes of some larvae allow them to eat the plant. Chrysanthemum Sesamin (inhibitor of detoxifying enzymes) protects plant from larvae again. istockphoto.com Plant produces pyrethrin (toxin). Fig 3.2: Chrysanthemum and the production of pyrethrin. plant communication Unbelievable as it may seem, plants are even able to communicate with each other for their mutual benefit. Chemicals are the way plants communicate with each other. Leaves damaged by pathogens or insect attack produce defensive molecules, such as ethene (ethylene), which can travel through the air to other plants to activate their plant defence genes. These volatile chemicals or ‘signalling’ molecules serve as a warning to others of the same species that they are under attack by herbivores. The neighbouring plants are then stimulated to produce their own chemical responses against the herbivores. When a willow tree (Fig 3.3) is attacked by insects, the leaves produce a chemical called salicin in response. This chemical travels through the tree and induces it to produce pathogenic chemicals (phytoalexins) that protect the tree from further attack. Some of the volatile salicin becomes airborne and is sensed by other willow trees nearby, warning them of the insect attack so that they can start to produce defensive chemicals too. © te ah o o t e k ur a p o un a m u istockphoto.com Acacia trees, for example, produce tannins to defend themselves when they are grazed on by animals such as giraffes. The tannins make the leaves unpalatable so after a few mouthfuls the giraffe moves away to another tree. But the acacias release the gaseous hormone ethene into the air. The airborne scent of the ethene is picked up by other acacia trees, which then start to produce tannins themselves as a protection from the grazers. Fig 3.3: Willow trees produce chemical-signalling molecules. BY3031 21 defence and cooperation Did you know? If you have ever taken an aspirin you will have used the willow’s own defences – aspirin is in fact a form of salicylic acid. Many cultures have used willow bark to lower fevers and ease pain. Studies in salicilin led to the development of aspirin, initially created from salicylic acid. 3A chemical defences Go to the Topic webpage to learn more about chemical defences and plant defence. carnivorous plants Plants are not always the victims of animal attacks. Sometimes, it is the plant that attacks. Plants that naturally grow in bogs lack essential minerals such as nitrogen, which is in low supply in boggy soils. These plants have overcome this deficiency by devouring insects and digesting their bodies to obtain the minerals they need. Some examples of carnivorous plants are the Venus fly trap, pitcher plants and sundews. Once the insect is inside the plant, digestive juices are released by the plant, digesting the prey and making its minerals available to the plant. 3B istockphoto.com Some of these plants, such as the Venus fly trap (Fig 3.4), have sensitive hairs that sense the presence of an insect when it lands on them and cause it to shut rapidly, trapping the insect. Pitcher plants are shaped in such a way that an insect entering the plant looking for nectar is unable to escape. Sundews have sticky hairs that trap insects. Fig 3.4: A Venus fly trap leaf showing its sensitive hairs. venus fly trap in action Go to the Topic webpage to watch videos of a Venus fly trap and a sundew catching flies. 22 BY3031 © te ah o o te k u ra p ou n a mu defence and cooperation plant–animal cooperation Plants not only repel animals in their environments. In many cases they cooperate successfully with animals for their mutual existence. Plants have evolved ingenious ways of getting pollinators to visit them and transfer pollen. Many animals may be plant pollinators, including bees and wasps, butterflies and moths, flies, some beetles and birds and even bats. The most obvious ways are through colourful petals and strong scents, but other flowers use more subtle ways to entice their pollinators to call. Some offer a reward in the form of nectar. Nectar is a sweet, sugary solution produced in special glands called nectaries, deep inside the flower. Some flowers such as pansies and foxgloves even have ‘honey guides’, which are markings such as spots, stripes or dark splotches that serve as guides to the centre of the flower where the nectaries lie. Did you know? Bees like white, yellow and blue flowers, and cannot see red very well. Bee-pollinated plants often have honey guides on them. Some honey guides are invisible to us but are visible to bees as they can see in the UV range of light. Bees can smell, so the flowers they pollinate are scented. Birds can see red and like bright-coloured flowers. tūī and mistletoe flower Go to the Topic webpage to watch a video of tūī opening mistletoe flowers. There are other plants that resort to mimicry to get animals to visit them and transfer pollen to other plants of the same species. A well-known example is that of the Australian hammer orchid (Drakaea species) (Fig 3.5) and thynnid wasps. The shape of the orchid flower resembles that of the female wasp, complete with shiny head and furry body. In addition the orchid releases a pheromone (a hormone that is volatile and travels through the air) to attract male wasps. When the male wasp tries to mate with the ‘female wasp’, it collects pollen from the orchid that is transferred to the next orchid the wasp visits. In this way, the orchid achieves pollination, but the wasp achieves nothing! © te ah o o t e k ur a p o un a m u wikimedia commons 3C Fig 3.5: The Australian hammer orchid resembles a female wasp. BY3031 23 defence and cooperation Some plants ensure their seeds are spread far from the parent tree to increase their chance of germination. They do this by having a close relationship with birds which eat the fruit containing the seeds. These then pass through the digestive tract of the bird and are deposited some distance from the parent tree. This mutualistic relationship ensures the birds have food from the tree and the tree increases the chance of its seeds successfully germinating. Some seeds actually require their tough seed coats to be scratched or given an abrasive treatment before they can germinate. This can occur naturally, as seeds are scratched by stones in the soil, but it also happens when the seeds pass through an animal’s digestive tract, as the digestive juices break down the seed coat. New Zealand’s native wood pigeon, the kererū, is a disperser of large fruits such as those of the karaka and taraire trees in our native forests, and is essential in aiding the regeneration of native forests. Some plants, such as acacias, have colonies of stinging ants living among their thorns. The ants feed on sugary nectar provided by the trees. They guard the trees and if another animal such as a browsing giraffe tries to eat the leaves, they swarm out and attack the browser. The relationship between the ants and the tree is a remarkable example of mutualism. The ants earn shelter and food, while the acacia gains protection from herbivores and competitors. 3D acacia and ants: mutualism Go to the Topic webpage to learn more about the mutualistic relationship between acacias and ants. plant defences Matagouri or wild Irishman (Discaria toumatou) is an example of a New Zealand native plant with thorns. These may be several centimetres long. Matagouri’s growth form is described as divarication (Fig 3.6). The plant grows up to six metres high and has very small leaves and small greenish-white flowers. It is widespread in the eastern South Island and Central Otago. doc 3E Fig 3.6: A matagouri plant. 24 BY3031 © te ah o o te k u ra p ou n a mu defence and cooperation Define the term ‘divarication’, and discuss why thorns and divarication are thought to be an adaptation to protect the plants against browsing by moa. Check your answers. © te ah o o t e k ur a p o un a m u BY3031 25 defence and cooperation 3F plant communication Lima beans (Phaseolus lunatus) may become infested with two-spotted spider mites (Tetranychus urticae) which damage their leaves. The lima beans produce volatile chemicals that can be detected by other bean plants growing nearby. When this distress signal reaches surrounding lima beans, they, too, begin to send the signal chemicals even though they are not under attack. In addition, the chemicals attract another species of spider mite that is carnivorous and preys on the two-spotted mite. Discuss how the production of volatile chemicals benefits the population of lima beans. Check your answers. 3G plant cooperation The two endemic mistletoe species (Peraxilla) both have large red showy flowers, unlike most New Zealand plant species, which have small and inconspicuous flowers. Both Peraxilla species have specialised ‘explosive’ flowers that require birds to tweak the buds before they will open. To open the flowers, birds grasp the top of the bud with their beaks and twist. This causes the flower petals to spring open (in less than a quarter of a second), and the bird can then insert its beak to drink nectar and in so doing pollinate the flower. Flower buds that are protected from all potential pollinators do not open their petals, although the petals do eventually separate as a unit from the base of the flower. The main birds that can twist open Peraxilla flowers are the tūī (Prosthemadera novaeseelandiae) and bellbird (Anthornis melanura), which are both endemic honeyeaters. 26 BY3031 © te ah o o te k u ra p ou n a mu defence and cooperation What is the term given to the relationship between the mistletoe and the birds? Discuss how this relationship is of benefit to both bird species and the mistletoe species. Check your answers. key words key points physical defences chemical defences plant communication plant cooperation •• Plants employ a range of physical and chemical defensive strategies to protect themselves from predation by herbivores. •• Plants can communicate with each other by chemical signalling. •• Plants cooperate with each other and with animals for their mutual benefit. © te ah o o t e k ur a p o un a m u BY3031 27 orientation responses learning intention In this lesson you will learn to: •• describe plant orientation responses to the abiotic environment. introduction Plants are acutely aware of their abiotic environment and are able to make suitable responses to stimuli. They are able to sense sunlight – not just its presence, but also the direction of the light, its intensity, and even the different wavelengths of light – and make appropriate responses to it. They can detect gravity, temperature, and the presence of water or chemicals, and can sense if they are touched. They can track the time of day and the time of year and adjust their growth accordingly. As almost no plants can move unaided, their responses are usually to alter their growth to move towards more favourable environments or away from less favourable environments. For example, if a plant is being shaded by a larger one, the smaller plant will grow taller or grow outwards to the side, to reach more sunlight. If a seed lands on the ground its roots will grow downwards and its stem upwards. In this lesson you will learn how plants respond to stimuli in their abiotic environment by orientating (positioning) themselves appropriately. tropisms A tropism is a growth response made by plants to external stimuli in their abiotic environment. The direction of the stimulus determines the direction of the growth response. Plants usually respond to stimuli such as sunlight, water, temperature, gravity, chemicals, touch and wind. The tropic response may be positive – the plant grows towards the stimulus; or negative – the plant grows away from the stimulus. Some tropisms you should know: •• light (phototropism): for example, a plant’s stem grows towards light so we say the stem is positively phototropic. A plant’s roots grow down into the soil, away from light, so the roots are negatively phototropic. The tree’s shoots grow towards the light, so they are positively phototropic. •• water (hydrotropism) •• temperature (thermotropism) •• gravity (geotropism or gravitropism) •• sun (heliotropism) •• chemicals (chemotropism) •• touch (thigmotropism) istockphoto.com 4 •• current (rheotropism). Fig 4.1: The tree’s roots grow down into the soil, away from light, so the roots are negatively phototropic. 28 BY3031 © te ah o o te k u ra p ou n a mu orientation responses nastic responses A nasty or nastic response is a non-directional movement of part of a plant in response to an external environmental stimulus. Nastic responses are made to the intensity of the stimulus rather than to the direction. For example, plants make nastic responses to the intensity of light, temperature and humidity, rather than to their direction. Nastic responses are turgor responses (turgor means the rigidity of the plant tissue due to inflation of the cells with water). You may have noticed that some flowers close at night and open in the daytime. These ‘sleep’ movements are in response to light intensity (photonasty) or temperature (thermonasty). These are nastic responses because they are triggered by external stimuli (light, temperature) but the response is directed internally. The movement is the result of growth or a turgor change. Crocus or tulip flowers close at night by the lower sides of the petals growing more rapidly and open by day by a more rapid growth of the upper sides. Some leaves show drooping or ‘sleep’ movements too. Several species of clover have structures called pulvini (singular: pulvinus). The pulvinus is a swelling at the base of the petiole or leaflet and has large parenchyma cells. Rapid turgor pressure changes in these cells cause the pulvinus to act as a hinge and bring about movement in the leaves. Movements that are triggered by the intensity of touch are called haptonastic. Haptonastic movements are perhaps the most fascinating of all plant movements – they bring about a rapid and elaborate response. Insectivorous plants snap shut in response to external pressure – like the Venus fly trap (which you learned about in the previous lesson) when its sensitive hairs are touched. Some of these movements are haptonastic. The mimosa plant (Mimosa pudica) responds very rapidly to touch. It is sometimes called the ‘sensitive plant’. If the leaflets at the tip are shocked by a sharp blow or even a sudden change in temperature or light intensity, they fold upwards in seconds. If the stimulus is sustained, successive pairs of leaflets fold upward and eventually the whole leaf drops. The stimulus is thought to be transmitted by a hormone that moves through the xylem; electrical changes are associated with its passage (but it does not have a nervous system). 4A mimosa leaves: haptonastic response Go to the Topic webpage to watch a video of mimosa leaves showing haptonastic responses. Note that tropisms are plant growth movements that are not usually reversible. Nastic movements on the other hand are reversible (for example, the crocus flower closes at night and opens again in the morning; the mimosa leaves close when touched and then open again). © te ah o o t e k ur a p o un a m u BY3031 29 orientation responses 4B plant responses 1. Define a tropism and a nastic response. Tropism: Nastic response: 2. Name each of the tropisms described below; say whether the tropism is positive or negative; and describe the biological advantage to the plants. a. A young stem grows upwards towards the light. Tropism: Biological advantage: b. Roots grow downwards into the soil. Tropism: 30 Biological advantage: BY3031 © te ah o o te k u ra p ou n a mu orientation responses c. Roots grow towards water. Tropism: Biological advantage: d. A stem grows away from the Earth’s gravitational pull. Tropism: Biological advantage: e. Tendrils (modified leaves) touch a trellis and coil around it. Tropism: Biological advantage: f. A climbing plant twines upwards around a supporting plant’s stem. Tropism: © te ah o o t e k ur a p o un a m u BY3031 31 orientation responses Biological advantage: g. Roots grow away from a copper pipe in the soil. Tropism: Biological advantage: h. A pollen tube grows from a pollen grain towards the chemicals given off by the flower’s ovary. Tropism: Biological advantage: 3. The leaves of Oxalis species droop at night. What is this plant response called, how does it occur and what is the biological advantage of this? Response: 32 How it occurs: BY3031 © te ah o o te k u ra p ou n a mu orientation responses Biological advantage: Check your answers. key words key points tropism nastic response •• A tropism is a growth response made by plants to external stimuli in their abiotic environment. The direction of the stimulus determines the direction of the growth response. The tropic response may be positive – the plant grows towards the stimulus; or negative – the plant grows away from the stimulus. •• A nasty or nastic response is a non-directional movement of part of a plant in response to an external environmental stimulus. Nastic responses are made to the intensity of the stimulus rather than to the direction. © te ah o o t e k ur a p o un a m u BY3031 33 control of plant growth 5 learning intention In this lesson you will learn to: •• discuss the plant hormones influencing plant growth. introduction You may have noticed that the leaves on a pot plant on a window sill turn to face the light. The leaves are displaying positive phototropism. If a seed lands on the soil, the roots will grow down into the soil and the stem will grow upwards. The roots are positively geotropic and the stem is positively phototropic. How do parts of the plant detect light or gravity or other stimuli? There are receptor cells in the leaves, shoots and roots that can detect the presence of the light and gravity stimuli and then respond appropriately to these stimuli. They do this by producing chemicals (hormones). Hormones are chemicals that are produced in one area of the plant and transported to other areas where they produce an appropriate growth response. plant hormones Charles Darwin and his son Francis carried out experiments on a species of Phalaris canariensis or canary grass. They sprouted the grass seeds and noticed that the coleoptile (a sheath that covers the first leaves as they emerge from the soil) bent towards the light coming from a window. The bending didn’t occur at the tip but in the elongating part behind the tip. Darwin covered the elongating part of the coleoptiles and shone light on the tips and the coleoptiles still bent towards the light. Next he covered the tips of the coleoptiles, shone light on them and found they did not bend. He concluded that there was some substance in the tip of the coleoptiles that was affected by light. This substance must be moving down the coleoptile and causing the lower part to bend towards the light. The light-sensitive region in the tip of the coleoptiles is called the receptor and the part below that elongates is the effector, where the response occurs. Many other scientists followed on from these experiments, and eventually the substance affecting growth was identified as a hormone and named ‘auxin’, from the Greek word ‘auxein’, which means to grow. Auxin was the first substance to be identified as a plant hormone. Auxin is in fact a group of hormones, the main one being indoleacetic acid (IAA). 5A auxins Go to the Topic webpage to learn about the early experiments to identify auxins. 34 BY3031 © te ah o o te k u ra p ou n a mu control of plant growth auxin Auxin is being made all the time by the cells in the tip of the shoot. The auxin diffuses downward from the tip into the rest of the shoot. Auxin makes the cells just behind the tip get longer. The more auxin there is, the faster they will grow. Without auxin they will not grow. When the light shines on a shoot from one side, the auxin at the tip concentrates on the shady side. This makes the cells on the shady side grow faster than those on the light side, so the shoot bends towards the light. This is explained in the diagram below: Auxins and phototropism Auxin is made here. Light Cells on this side grow quickly. Cells on this side grow slowly. Light The uneven concentration of auxin causes the shady side to grow faster than the light side, so the shoot bends towards the light. Auxin made in this tip diffuses unevenly down the shoot, concentrating on the shady side. auxin has opposite effects on stems and roots It seems from experimental evidence that the concentration of auxin has opposing effects on stems and roots. Roots are much more sensitive to auxin concentration than stems. Root growth is actually inhibited by high concentrations of auxin. Accumulation of auxin stimulates growth on this side of plumule. Accumulation of auxin is present on lower side of elongating zones of plumule and radicle. © te ah o o t e k ur a p o un a m u gravity Accumulation of auxin inhibits growth on this side of radicle. BY3031 35 control of plant growth Auxin accumulates by gravity on the lower sides of stems and roots. In the root (radicle), this high concentration inhibits growth on the lower side of the elongating cells so that the elongating cells on the upper side grow downward. In the stem (plumule), the high concentration of auxin on the lower side has the opposite effect, stimulating growth of the lower cells and making the stem bend upwards. Geotropism enables the roots to grow downwards in the soil irrespective of the seed’s orientation under the soil surface. This will help to anchor the plant and to absorb the water and minerals needed from the soil. But how does the seed tell up from down? It uses gravity receptors. These are small starch grains (called amyloplasts or ‘statoliths’) in the cells. They move to the lower side of the cells and affect the distribution of auxin so that there is more auxin on the underside. Positive geotropism in roots Whichever way up a seed is planted, its radicle always grows downwards. If you have an old pot plant, you could try a similar experiment by simply lying the potted plant on its side and seeing what happens to it over a two-week period. Potted plant lying on its side Hormone accumulates on lower side of stem and stimulates cell growth – stem turns up. 36 BY3031 Hormone accumulates on lower side of root and inhibits cell growth – root turns downwards. © te ah o o te k u ra p ou n a mu control of plant growth apical dominance The tip (apex) of a plant produces the most auxin, so the plant grows upwards. The side shoots (laterals) do not grow, as auxin inhibits their growth. However, if the tip of the shoot is cut off (a process called ‘pinching out’ by gardeners) then the side shoots will develop and the plant will grow bushier! effects of plant hormones There are five major groups of plant hormones involved in plant growth. These are: •• auxins •• gibberellins •• cytokinins •• abscisic acid •• ethene (ethylene) gas. The main effects of these plant hormones are summarised in the table below. Their effects are more complex and interactive than given here. Note that you do not need to remember the specific effects of these hormones. You do, however, need to understand that they are produced in response to environmental stimuli and they allow plants to make appropriate growth-tropic responses, to help them adapt and survive changing environmental conditions. main plant hormones and their effects Hormone Main effects Auxin Promotes stem elongation. Controls cell enlargement. Causes apical dominance by suppressing growth of lateral buds. Stimulates cell division in cambium (secondary vascular growth). Stimulates root initiation. Suppresses root elongation. Delays onset of leaf fall. Delays fruit ripening. Stimulates growth of flower parts. Gibberellin Promotes stem elongation. Delays dormancy and leaf fall. Breaks dormancy in seeds and buds. Cytokinin Stimulates cell division. Promotes growth of young fruit. Balances root and shoot growth. © te ah o o t e k ur a p o un a m u BY3031 37 control of plant growth 5B Abscisic acid Inhibits growth, produces winter dormancy. Induces fruit fall. Promotes seed dormancy. Acts on guard cells; produced in response to water stress and promotes closing of stomata. (The effects of abscisic acid are generally the opposite of those of auxins, gibberellins and cytokinins.) Ethene (ethylene) gas Induces fruit to ripen. Aids leaf fall. plant hormones Go to the Topic webpage to read an overview on plant hormones. 5C plant hormones 1. When water levels in a plant are low, due to low rainfall or drought conditions, the synthesis of abscisic acid in the plant is stimulated. One of the functions of abscisic acid is to stimulate the closing of the pairs of guard cells surrounding stomata in the leaves. Discuss how this response helps the plant to survive the dry conditions. 38 BY3031 © te ah o o te k u ra p ou n a mu control of plant growth 2. A seedling is placed horizontally on the soil surface. Explain the role of auxin in the geotropic (gravitropic) response of both the shoot and the root of this seedling and discuss why this response helps the seedling to survive and grow into a plant. Check your answers. key words key points plant hormones receptor effector •• Hormones are chemicals that are produced in one area of the plant and transported to other areas where they produce an appropriate growth response. •• Receptor cells in the leaves, shoots and roots detect the presence of the light and gravity stimuli and then respond appropriately to these stimuli by producing hormones. •• The effector is the region of the plant where the growth response occurs. © te ah o o t e k ur a p o un a m u BY3031 39 timing responses learning intentions In this lesson you will learn to: •• explain plant photoperiod timing responses •• explain the plant phytochrome system. introduction If you have a garden or live on a farm you have probably noticed seasonal changes in plant growth and development. Most plants flower in the heat of summer; spring is a good time to plant seedlings; many fruit and vegetables ripen in autumn; some leaves change colour and fall from deciduous trees in autumn; other plants go dormant in winter; and new leaves and buds appear in spring. If you are observant you may have noticed that some flowers open in the daylight and close at night while others, like the evening primrose (Oenothera biennis) (Fig 6.1), open in the late afternoon. Some flowers are scented and produce nectar in the daytime, and others open and are scented at night. Obviously, plants are able to detect environmental stimuli that tell them the time of day and the time of year, so that they can adjust their growth and flowering and fruiting times accordingly. timing rhythms istockphoto.com 6 Fig 6.1: The evening primrose flowers open in the evening, when light levels are lower. Some plants such as the evening primrose above show daily (or diurnal) rhythms that may be triggered by light or temperature. Daily rhythms are also called circadian rhythms (circadian means ‘about a day’, from the Latin ‘circa’ meaning ‘about’ and ‘dies’ meaning ‘a day’). Flowers that open in the sunlight and close when the sun sets are displaying a circadian rhythm. Leaves that droop at night and are upright during the day are also displaying a circadian rhythm. Some plants show yearly (circannual) rhythms, as they produce buds and new leaves in spring, flower at one time of the year (often spring or summer), lose their leaves in autumn, and go dormant at other times (usually in winter, but in summer for some plants). These plants are following a yearly cycle of growth and development. •• Annual plants grow, flower and set seed during one year (for example, petunias and pansies). •• Biennial plants grow leaves and store food in their first year and flower in the second year (for example, carrots and foxgloves). •• Perennials grow and flower every year for a number of years. Many die down and become dormant, usually over winter, then grow new leaves in spring (for example, dahlias and lavender). •• Ephemerals are desert plants that germinate, grow, flower and set seed in a few weeks, after rainfall. They contain a chemical that inhibits the seeds from germinating until sufficient rain has fallen. Then they carry out their life cycle quickly, before the desert dries out again. 40 BY3031 © te ah o o te k u ra p ou n a mu timing responses If the stimulus that brings about these plant responses comes from outside the plants, it is called ‘exogenous’ (‘exo’ means ‘outside’). A stimulus may also be produced internally within the plant. This is termed ‘endogenous’ (‘endo’ means ‘inside’). change in leaf colour change from leaf development to flower development grass tillering leaf drop in deciduous trees in autumn plant responses to day length bud break of fruit tree in spring tuber and bulb formation start of winter dormancy germination of some seeds, for example, lettuce photoperiod and phytochrome The control over the life cycle of plants (from seed → reproductive maturity → seed) is achieved by the cyclic change in day length (and temperature). The further plants are from the equator, where days are almost a constant 12 hours long, the greater the seasonal variation in day length. In plants, phenomena such as flowering, fruit and seed production, bud and seed dormancy, leaf fall, and germination are closely attuned to seasonal differences such as day length and temperature. The survival of the plant depends on these. The process that involves the most profound change is flowering, when shoot meristems (growing points) switch from producing leaves and lateral buds to producing flowers. It is vital for plants to coordinate their flowering times, so that pollination can take place. The flowers of the same species need to flower in synchronisation so that pollen can be transferred from one to another, and at the right time of year for their pollinators such as insects and birds to be present. © te ah o o t e k ur a p o un a m u BY3031 41 timing responses There are many environmental cues that affect flowering, such as temperature, but it is mainly the length and sequence of the light and dark periods, rather than the total amount of light received, that stimulates reproduction (flowering). The term photoperiodism is used to describe the plants’ biological response to the relative amounts of light and darkness in any 24-hour period (the photoperiod). Plants use the photoperiod to tell the length of day and therefore which season it is. How can plants sense the length of day and night? They use a sensory blue-green pigment, called a phytochrome. Phytochromes are protein pigments in plants that are sensitive to red and infrared light. Two interchangable forms exist: •• Pr, which absorbs red light and •• Pfr, which absorbs infrared light. Each form changes back to the other when it absorbs light. On absorbing light in the daytime, Pr converts to Pfr. At night, Pfr reverts back to Pr. Pfr is the biologically active form that acts as a switch to turn on plant responses. This is shown in the diagram below: in daylight Pr Pfr plant response at night inactive form active form Phytochromes are important in the control of flowering. They also act in: •• the germination of seeds, which need a brief exposure to light before they germinate •• stem elongation (infrared light) •• leaf expansion (red light) •• growth of side roots (infrared light). The photoperiod stimulus is sensed in the leaves via the phytochrome system. A chemical messenger is then sent to the buds to form flowers. (The messenger has been called ‘florigen’, but has not yet been isolated and may be a mixture of hormones.) 6A phytochromes Go to the Topic webpage to learn more about the phytochrome system and the control of flowering. 42 BY3031 © te ah o o te k u ra p ou n a mu timing responses photoperiod and flowering reponses From the early twentieth century, scientists discovered the importance of the photoperiod in flower formation. It was shown that tobacco plants would flower only after exposure to a series of short days. This occurred naturally in autumn, but could also be induced artificially in a greenhouse by limiting daylight to seven hours. Plants that need a short day and long night to flower are called short-day plants. They will only flower if the photoperiod is less than a certain critical length. Other plants require long days and short nights for flowering (long-day plants). In order to flower, these plants need a photoperiod that exceeds a certain critical length. Some other plants flower when they are mature, whatever the photoperiod is. These are dayneutral plants. Later advances to our understanding have shown that it is actually the length of the dark period that is the critical factor. So short-day plants are really long-night plants. If these are grown in short days but the long night is interrupted by a short light period, then flowering is prevented. Similarly, long-day plants (short-night plants) will flower in short days if the long night is interrupted. Long-day plants will flower only when red light or a long period of sunlight causes an accumulation of Pfr. Short-day plants flower when red light or a long period of darkness causes an accumulation of Pr. Short-day plants (SDP) typically flower in the spring or autumn when the length of day is short. Long-day plants (LDP) typically flower during the summer months of longer photoperiod. This is explained in the following diagram. = flowering Short-day plant (SDP) = no flowering Long-day plant (LDP) 24 hours 16 hours light (long day) 8 hours light (short day) 8 hours dark 16 hours dark 8 hours light 6B 8 hours light red 8 hours light red infrared long-day/short-day plants Go to the Topic webpage to view an animation explaining long-day/short-day plants. © te ah o o t e k ur a p o un a m u BY3031 43 timing responses more plant responses to the abiotic environment temperature (vernalisation) Temperature changes can act as cues for plant responses. Some plant species require exposure of a growth stage to cold before flowering can proceed. This need for a cold spell is called vernalisation. Here, the stimulus is picked up by the mature stem apex or by the embryo of the seed, not the leaves, as in photoperiodism. Long-day plants (such as cabbage), short-day plants (chrysanthemum) and day-neutral plants (ragwort) can all require vernalisation. The length of chilling can vary from four days to three months, with temperatures around 4°C generally being most effective. During vernalisation, the level of gibberellins increases. Gibberellins can be used artificially on unvernalised plants to substitute for vernalisation. Seeds may also be subjected to cold by placing them in a refrigerator if they are to be grown out of season. Some species have their seeds planted in the autumn so that they germinate before the winter cold because flowering will not occur unless the young seedlings have been exposed to cold. Plants from the southern areas of New Zealand may not survive when planted in more northerly parts of the country, because they may not be exposed to sufficiently cold periods for a long enough time. importance in plant survival Both photoperiodism and vernalisation work to synchronise the reproductive behaviour of plants with their environment to ensure reproduction at the most favourable times of the year. This will also ensure that members of the same species will flower at the same time, encouraging crosspollination and cross-fertilisation, with the advantages these bring in genetic variability. dormancy in plants Dormancy occurs when growth and development in plants cease and the plants’ metabolic rates fall to a point which is only just sufficient to keep the cells alive. This is the normal winter state of most plants in temperate regions. Dormancy is brought about by light and temperature acting through hormones. Winter buds in temperate trees (such as birch, beech and sycamore) form as a photoperiodic response to the shortening days in autumn. This stimulus is perceived in the leaves and hormones are produced, which leads to a build-up in the chemical abscisic acid. Abscisic acid inhibits growth and also induces leaf fall. Storage organs (for example, bulbs and tubers) are also involved when the plant goes into a dormant state, and again, photoperiodism is involved. Short days induce tuber formation in potatoes and long days induce onion bulb formation. 44 BY3031 © te ah o o te k u ra p ou n a mu timing responses leaf fall (abscission) Leaf fall can result from a number of factors. These include: •• water shortage as a result of drought conditions or the freezing of water in the soil •• lowering of light intensity •• a drop in temperature •• shortening of day length. As a result of a build-up in abscisic acid, a layer of weakened cells collects at the base of the petiole. The leaf eventually breaks off from the parent plant at this point. This is most obvious in deciduous plants. rhythm and phase Leaves of the plant Bryophyllum (a common New Zealand weed, also called ‘mother of millions’) show a circadian rhythm in their output of carbon dioxide gas. This rhythm can be altered by a number of abiotic factors, including light and temperature. Figure 6.2 shows the normal rhythm of the Bryophyllum in continuous darkness at constant temperature, and the altered rhythm when the Bryophyllum was exposed to four hours of light (dark line on the graph). output 60 CO₂ exchange (relative amount) Continuous darkness, at constant temperature 4 hours light 40 leaf in darkness (normal rhythm) leaf in darkness plus 4 hours light 20 0 -20 -40 -60 -80 -100 -120 -140 intake 0 midnight 0 0 0 0 midnight Time of day NZQA 6C Fig 6.2: Rhythm of CO2 production in continuous darkness and with 4 hours of light, once around midnight. © te ah o o t e k ur a p o un a m u BY3031 45 timing responses Figure 6.3 shows how the normal rhythm in continuous light at 15°C can be altered (dark line) by increasing the temperature to 40°C for three hours without altering the light. output 60 CO₂ exchange (relative amount) 40 Continuous light at 15°C untreated leaf (normal rhythm) 3 hours at 40°C 20 altered rhythm plus 3 hours of higher temperature 0 -20 -40 -60 -80 -100 -120 0 0 midnight midnight 0 0 0 Time of day NZQA -140 intake Fig 6.3: Rhythm of CO2 production in continuous light except for 3 hours of higher temperature. Both figures show the rhythm over five days. 1. What does the term ‘circadian’ mean? 2. Explain how the exposure to four hours of light has altered the circadian rhythm (Fig 6.2). 46 BY3031 © te ah o o te k u ra p ou n a mu timing responses 3. How is the effect on the circadian rhythm of the exposure to 40°C (Fig 6.3) different to the effect of the exposure to light (Fig 6.2)? Assume that both the differences in duration of ‘treatments’ (4 hours of light versus 3 hours of higher temperature) and timing of treatments (3 hours leading up to midnight versus 2 hours on either side of midnight) can be ignored. 4. Explain from the data why this rhythm is endogenous. Check your answers. 6D photoperiod and flowering The changing length of day and night regulates the onset of flowering in many plants. 1. Give the term that describes the regulation of activity in plants by day and night length. 2. Describe two advantages of having the time of flowering regulated by an environmental factor. © te ah o o t e k ur a p o un a m u BY3031 47 timing responses 3. Name one process in plants, other than flowering, controlled by the length of day and night. 4. Study the information in the diagram below, and then answer the questions that follow. Equal length days and nights Short days, long nights flower Cocklebur Corn flower Long days, short nights flower NZQA Spinach flower a. Is spinach a short-day or a long-day or a day-neutral plant? 48 BY3031 © te ah o o te k u ra p ou n a mu timing responses b. Describe the environmental factor that initiates flowering in short-day plants. c. Explain how cocklebur, a short-day plant, can flower when day and night lengths are equal. d. Poinsettia is a short-day plant with a critical day length of around 13 hours. Poinsettia plants were put in a glasshouse under conditions of short days and long nights. Explain what will happen to the flowering if the day period is interrupted by a few minutes of darkness. Check your answers. key words key points circadian (diurnal) rhythm endogenous exogenous photoperiod photoperiodism phytochrome short-day plant (SDP) long-day plant (LDP) vernalisation dormancy abscission •• Stimuli may be endogenous (internal) or exogenous (external) to a plant. © te ah o o t e k ur a p o un a m u •• Plant responses may follow a circadian rhythm. •• Photoperiodism describes the plants’ biological response to the relative amounts of light and darkness in any 24hour period (the photoperiod). •• Plants sense the photoperiod using phytochromes. BY3031 49 7 teacher-marked assignment learning intention In this lesson you will: •• review your progress over this topic and practise exam-type questions. introduction In this lesson, have a quick look back at all the lessons you have completed in this topic. Think about what you have learned. When you are ready, try the teacher-marked assignment BY3031A. If you did not receive this with your booklet, contact your teacher. When you have finished, complete the self-assessment section in the teacher-marked assignment. Send the self-assessment and the teacher-marked assignment to your teacher. Make sure that you have written your name and ID number on the cover sheet of the teachermarked assignment. You can also use a label if you have one at hand. By post: Put the self-assessment and teacher-marked assignment in the plastic envelope provided. Make sure that the address card shows the address for Te Aho o Te Kura Pounamu (The Correspondence School). Seal the envelope with tape before you post it. By email: Scan the pages including the cover sheet and email to your teacher. The standard format for Te Kura teacher email addresses is: [email protected] If you aren’t sure who your teacher is, call 0800 65 99 88. Before you finish off this topic you should have agreed your next steps with your teacher. If you do not have your next set of study materials, contact your teacher immediately. If you are not sure what to do next, ask your teacher for advice. 50 BY3031 © te ah o o te k u ra p ou n a mu answer guide 8 1. the environment: an overview 1C Stimulus: a change in the environment that causes a response in an organism Habitat: the place where an organism lives Biotic environment: the influences of living organisms on one another Abiotic environment: the non-living influences on an organism Ecosystem: a community of living organisms (biotic environment) together with non-living factors (abiotic environment) and the nutrient cycles and energy flows that connect all Intraspecific: relationships within a species Interspecific: relationships between species Adaptation: an inherited characteristic that enables an organism to survive and reproduce in its habitat. Adaptations may be structural, behavioural or physiological Tolerance: an organism’s ability to survive variation in its environmental conditions Acclimatisation: the ability of an organism to adjust its tolerance limits or optimum range to slow changes in its environment Ecological niche: a description of how the organism feeds, the opportunities provided by the habitat and the adaptive features of the organism which allow it to take advantage of these opportunities © te ah o o t e k ur a p o un a m u BY3031 51 answer guide 1D Activity 1.2: Adaptations Answers will vary but may include: Achieved answer Merit answer Gives at least two adaptations. Structural adaptations include: Links the adaptations to the falcon’s niche: •• wings for flight •• The falcon’s niche is that of a bird of prey. Its wings allow it to fly long distances to seek prey. •• sharp talons to catch and hold onto prey •• excellent eyesight •• strong beak to kill prey. •• Its sharp eyesight helps it locate prey even while flying at speed. •• The falcon can catch prey while in flight and uses its strong talons to hold onto its prey and its strong beak to kill its prey. These adaptations make the falcon a successful hunter. 2. competition and cooperation 2F 1. Achieved answer Merit answer Defines ‘allelopathy’. As for Achieved AND explains competitive advantage. For example: Allelopathy is the production of toxic chemicals in the roots and leaves of a plant that prohibits the growth of neighbouring plants. 52 BY3031 •• This is an advantage to the plant producing the chemicals because it stops or greatly reduces the growth of neighbouring plants which would compete with the plant for resources such as sunlight, water, minerals from the soil, space to grow, availability of pollinators, etc. This gives the allelopathic plant a competitive edge in surviving in the environment, to the detriment of its neighbours. © te ah o o te k u ra p ou n a mu answer guide 2. 2G Achieved answer Merit answer Epiphytes grow high in the branches of canopy trees so they are able to access more sunlight than plants growing lower in the forest. As for Achieved AND explains competitive advantage. This gives them an advantage as they can carry out more photosynthesis and therefore grow bigger and quicker than lower growing plants that receive less sunlight. Being epiphytic also protects these plants from ground-feeding herbivores. Achieved answer Merit answer Excellence answer For example: The fungal hyphae of the Tuber melanosporum extend from the host tree into the surrounding soil. This increases the surface area for absorption of water and nutrients by the fungus or the host tree. As for Achieved AND explains the advantage. As for Merit AND gives detailed full comparison/ answer. © te ah o o t e k ur a p o un a m u For example: The fungi also secrete growth factors that stimulate the roots to grow and branch and may produce antibiotics that may protect the tree from pathogens. In return the fungus receives carbohydrates made by photosynthesis in the tree’s leaves. For example: Because of the extra root area for absorption of water and minerals and protection that the Tuber melanosporum provides, trees infected with this fungus grow better than other trees that are not infected. BY3031 53 answer guide 3. defence and cooperation 3E Achieved answer Merit answer Excellence answer Definition given. As for Achieved AND explains advantage of either divarication OR thorns. As for Merit AND gives full account/comparison of advantage of both thorns and divarication. For example: Divarication makes it difficult for moa to reach the inner, larger leaves. For example: It is thought that the many divaricating species in NZ arose as an adaptation to browsing by moa, as the combination of thorns and divarication means it is difficult for browsers to reach the inner, larger leaves. The shrub is more tightly bound in a compact shape, making it harder to break off a branch. The extra toughness of the stems protects the plant against browsing. The high proportion of stem to leaf also made the plant less attractive to moa. For example: Divarication is a form of growth where plants branch repeatedly and produce a tangle of stems with the outer branches having smaller and fewer leaves than the inner branches. The stems of divaricating plants are often tough and difficult to break. 54 BY3031 © te ah o o te k u ra p ou n a mu answer guide 3F Achieved answer Merit answer Excellence answer Describes at least one effect of the production of volatile chemicals. As for Achieved AND explains the effect. As for Merit AND gives full account/comparison linking several effects. For example: The production of volatile chemicals by the lima bean has two effects. Firstly, the chemicals benefit the plants growing nearby by warning them of the attack by the two-spotted spider mites. 3G For example: These plants can then produce chemicals of their own to repel any spider mite attacks. For example: In addition, the chemicals attract a different species of spider mite that does not eat the lima beans but preys on the twospotted spider mites. This reduces the numbers of two-spotted mites eating the lima beans, thus benefiting the plants. Achieved answer Merit answer Excellence answer States the type of interrelationship. As for Achieved AND explains benefit. For example: The relationship between the bird species and the mistletoe species is mutualism (symbiosis), as both the bird species and the mistletoe species benefit. For example: The birds benefit from opening the mistletoe flowers by obtaining nectar, which is a nutritious food source. As for Merit AND gives full account/comparison of benefit for all three species. OR The flowers benefit from being opened by the birds by being pollinated as the birds carry pollen from the open flower to the next flower that they open. © te ah o o t e k ur a p o un a m u For example: Cross-pollination could benefit the mistletoe population by creating genetic variation in the mistletoe population/ gene pool and, of course, it may increase seed production. Having access to this energy-rich nectar may enable birds to produce more eggs, resulting in more offspring surviving. This in turn could increase their population size. BY3031 55 answer guide 4. orientation responses 4B Note: all these answers are at Achieved level. 1. A tropism is a growth response made by plants to external stimuli in their abiotic environment. The direction of the stimulus determines the direction of the growth response. Tropisms may be positive or negative. A nasty or nastic response is a non-directional movement of part of a plant in response to an external environmental stimulus. Nastic responses are made to the intensity of the stimulus rather than to the direction. 2. a. A young stem grows upwards towards the light. Tropism: Positive phototropism Biological advantage: The stem and leaves reach sunlight that is needed for photosynthesis. b. Roots grow downwards into the soil. Tropism: Positive geotropism (or gravitropism) Biological advantage: The roots anchor the plant in the soil and the roots can find water and minerals. c. Roots grow towards water. Tropism: Positive hydrotropism Biological advantage: The roots can find water, which is essential for all chemical processes in the plant. d. A stem grows away from the Earth’s gravitational pull. Tropism: Negative geotropism (or gravitropism) Biological advantage: The stem does not grow into the soil where it will not be able to get sunlight for photosynthesis. e. Tendrils (modified leaves) touch a trellis and coil around it. Tropism: Positive thigmotropism Biological advantage: The plant can be supported as it grows. f. A climbing plant twines upwards around a supporting plant’s stem. Tropism: Positive phototropism Biological advantage: The plant can be supported as it grows. The plant can grow upwards to reach the sunlight, for photosynthesis. Flowers can be held up, so that pollinators can reach them. 56 BY3031 © te ah o o te k u ra p ou n a mu answer guide g. Roots grow away from a copper pipe in the soil. Tropism: Negative chemotropism Biological advantage: The roots avoid a potentially toxic substance that may harm the plant’s growth. h. A pollen tube grows from a pollen grain towards the chemicals given off by the flower’s ovary. Tropism: Positive chemotropism Biological advantage: The pollen tube will grow towards the ovule in the ovary and the plant will be pollinated so that seeds can form to produce new plants. 3. Response: The leaves of Oxalis species drooping at night is a photonastic response. How it occurs: This response is brought about by changes in turgor pressure in the cells at the base of the leaves. Biological advantage: The drooping of leaves reduces heat and water loss during the night and reduces the amount of moonlight on the leaves (so that the plant does not photosynthesise at night). © te ah o o t e k ur a p o un a m u BY3031 57 answer guide 5. control of plant growth 5C 1. 58 Achieved answer Merit answer Excellence answer Describes how stomata open and close. As for Achieved AND explains role of stomata. For example: The paired guard cells on either side of the stomata open and close the stomata by changes in their turgor (water) pressure. For example: The stomata control gases and water vapour entering and exiting the plant. The plant has to balance the amount of carbon dioxide coming into the plant for photosynthesis, and the amount of water vapour escaping through the open stomatal pores by transpiration. As for Merit AND detailed answer given including role of water in life of plant. BY3031 For example: By producing abscisic acid the plant is responding to the environmental stimulus of lack of water. The abscisic acid promotes the closing of the stomata, which prevents water vapour from leaving the plant by transpiration, thereby regulating water loss and conserving the water available to the plant. Plants need water for all their metabolic processes. © te ah o o te k u ra p ou n a mu answer guide 2. Achieved answer Merit answer Excellence answer Defines geotropic (gravitropic). As for Achieved AND states the effect of auxin on shoot/root. As for Merit AND full answer given, explaining the effect of auxin on shoot AND root. For example: The geotropic responses of both shoots and roots are due to the presence of auxin. Shoots grow towards light and away from the force of gravity, so shoots are positively phototropic and negatively geotropic. Roots grow towards gravity, so they are positively geotropic. © te ah o o t e k ur a p o un a m u For example: When a developing shoot is placed horizontally, auxin moves to the lower side of both the shoot and the roots. The growth rate of the cells on the upper and lower parts of both the shoot and root differ in response to the presence of auxin. For example: In the shoots auxin causes elongation of the cells on the lower surface, which causes the shoots to grow upwards. Root elongation on the other hand is inhibited by high levels of auxin on the lower side of the root. The cells that elongate the most are therefore found on the upper surface of the root, so the root grows downwards. These tropic responses are essential for the seedling’s survival as they ensure the shoots grow upwards towards the light, which is needed for photosynthesis, and the roots grow downwards to anchor the plant in the soil and absorb water and nutrients needed for plant growth. BY3031 59 answer guide 6. timing responses Achieved answer 6C Merit answer 1. Circadian means daily/about a day. 2. It has shifted the rhythm by 12 hours/ half a cycle. As for Achieved AND links this shift to CO2 production. For example: •• Because of the one-off treatment of four hours of light around midnight, the peak of CO2 production shifts to being at midnight for at least the next three days, instead of being at midday. 3. Gives one difference. For example: •• The direction of the phase shift in one direction is opposite to the direction of the phase shift in the other. •• While the rhythm peak reduces with time in Fig 6.3 this peak remains the same for Fig 6.2. •• While the period (time between two peaks) is increasing in Fig 6.3, this period remains the same in Fig 6.2. 4. The rhythm is endogenous because it is maintained in the absence of external cues/under constant environmental conditions. 60 BY3031 As for Achieved AND gives evidence of rhythm being endogenous. For example: •• Both figures maintain a cyclic pattern of CO2 production with highs and lows appearing even when there is no change to the environmental conditions/ treatment. © te ah o o te k u ra p ou n a mu answer guide 6D 1. Photoperiodism (not photoperiod) •• ensures flowering at a time when pollinators are available •• flowers in suitable environmental conditions for seed production or germination •• time of flowering coordinated for plants of the same species to ensure pollination. •• leaf fall •• bud dormancy •• bulb and tuber formation •• chlorophyll synthesis •• dormancy •• vernalisation •• germination 2. a. Long-day plant. b. When the length of the night exceeds a certain period. c. Achieved answer Merit answer Correct answer given. As for Achieved AND states number of hours for critical day length. For example: Cocklebur, being a short-day plant, will flower: •• when the night length that triggers flowering is more than or equal to 12 hours For example: Cocklebur has a critical day length (CDL) of 12 hours or less. •• when day and night length are equal •• nights are getting longer. d. Achieved answer Merit answer Correct answer given. As for Achieved AND gives explanation. The poinsettia: For example: Because length of darkness, not length of day, triggers flowering. •• will flower •• nothing will change •• no effect on flowering. © te ah o o t e k ur a p o un a m u BY3031 61 glossary abscission Leaf fall from deciduous trees in autumn. abiotic environment The non-living environment; the non-living influences on an organism. acclimatisation The ability of an organism to adjust its tolerance limits or optimum range to slow changes in the environment. adaptation An inherited characteristic that enables an organism to survive and reproduce in its habitat. Adaptions may be structural, behavioural or physiological. allelopathy The production of toxic chemicals in the roots and leaves of a plant that prohibit the growth of other plants. auxin A plant hormone that affects growth and development. biotic environment The living environment; the influences of living organisms on one another. circadian An event repeating itself about every 24 hours. diurnal Daily or circadian rhythm. dormancy A resting phase in an organism when metabolic rate falls to a low level. ecological niche A description of how the organism feeds, opportunities provided by the habitat and the adaptative features of the organism that allow it to take advantage of these opportunities. ecosystem A community of living organisms (biotic environment) together with non-living factors (abiotic environment), and the nutrient cycles and energy flows that connect all. endogenous A factor or stimulus that originates within an organism. exogenous A factor or stimulus that originates outside an organism. habitat The place where an organism lives. long-day plants Plants which flower in response to a day length longer than a critical period. nastic response A non-directional movement of part of a plant in response to an external environmental stimulus. periodicity Regularly changing behaviour. 62 BY3031 © te ah o o te k u ra p ou n a mu glossary photoperiod The relative amount of light and darkness in a 24-hour period. photoperiodism A changing pattern or response based on length of light available. phytochrome A protein pigment in plants sensitive to red and infrared light. plant hormone A chemical that is produced in one part of the plant and has effects in another part. short-day plants Plants that flower in response to a day length shorter than a critical period. stimulus A change in the environment that causes a response in an organism. symbiotic (mutualistic) association An association between two organisms for their mutual benefit. tolerance An organism’s ability to survive variation in its environmental conditions. tropism A growth response made by plants to external stimuli in their abiotic environment. vernalisation The exposure of a plant to a period of low temperature to promote later growth and flowering. © te ah o o t e k ur a p o un a m u BY3031 63 acknowledgements Every effort has been made to acknowledge and contact copyright holders. Te Aho o Te Kura Pounamu apologies for any omissions and welcomes more accurate information. Photos Cover: Korukoru, Scarlet mistletoe in flower (Peraxilla [Elytranthe] colenso) on beech tree, # 11441FP13 © Rob Suisted © naturespic.co.nz. Sunset friends, # 17889609 © Alex Koch © istockphoto.com. Calico kitten playing with a toy, # 13048177 © Erik Zunec © istockphoto.com. Kalahari meerkats, # 16440743 © Peter Malsbury © istockphoto.com. Cape cobra, # 1694906 © Nico Smit © istockphoto.com. New Zealand falcon, #39587CE00 © Rob Suisted/www.naturespic.com, Wellington, NZ. Used by permission. Tree tops, © Don Laing, Wellington, NZ: Te Aho o Te Kura Pounamu. Used by permission. Gorse, Close up of gorse flower 1, © Don Laing, Wellington, NZ: Te Aho o Te Kura Pounamu. Used by permission. Walnut, Ready to fall, # 14430606 © Barry Sutton © istockphoto.com. Flowering Northern rātā trees in forest canopy (Metrosideros robusta), Kahurangi National Park, West Coast, NZ, # 6548FG@01 © Rob Suisted © naturespic.co.nz. Hemiparasitic red mistletoe (Peraxilla [Elytranthe] tetrapetala), NZ native bush on silver beech tree (Nothofagus menziesii) Arthur’s Pass National Park, NZ, # 5277FP06 © Rob Suisted © naturespic.co.nz. Feijoa flower, # 4956437 © Alan Drummond © istockphoto.com. Marigolds, # 16448303 © Waseef © istockphoto.com (used twice). Plant root showing nodules; Lichen, by Veronica Armstrong, © Te Aho o Te Kura Pounamu. Used by permission Lichens growing on a tree, © Don Laing, Wellington, NZ: Te Aho o Te Kura Pounamu. Used by permission. Perching lilies, photo by Phil Bendle, retrieved from http://www.terrain.net.nz/friends-of-te-henui-group/new-plant-page/perchinglily.html, © 2008–2012 T.E.R:R.A.I.N. Used in any medium for education and its promotion by permission. Coprosma rhamnoides, by Rudolph89, Wikimedia Commons, retrieved from http://commons.wikimedia.org/wiki/File:Coprosma_ rhamnoides_11.JPG, 9 November 2012. Creative Commons CC0 1.0 Universal public domain dedication. Chryanthemum, # 19377214 © Ivan Vasilev © istockphoto.com (used twice). Chryanthemum leaves, # 17115637 © Thomas Acop © istockphoto.com. Green caterpillar, # 13980701 © Alasdair Thomson © istockphoto.com. Willow at riverside, # 12454245 © Jose Ignacio Soto © istockphoto.com (used twice). Venus fly trap, # 7122250 © Mark Goddard © istockphoto.com. Sketch of Drakaea elastica (Hammer Orchid) from John Lindley’s A Sketch of the Vegetarian of the Swan River Colony, http:// commons.wikimedia.org/wiki/File:A_Sketch_of_the Vegetation_ofthe_Swan_River_Colony_–_Figure_3.png, accessed September 2012. Matagouri, Discaria toumatou, close up of flowers and spikes, © Chris Rance © DOC. Herb – Oenothera biennis, # 15609495 © Davidenko Andrey © istockphoto.com. 64 BY3031 © te ah o o te k u ra p ou n a mu acknowledgements Graphs Circadian rhythm of Bryophyllum in 1) darkness compared to light; 2) light compared to heat, source not given; from University Entrance, Bursaries and Scholarships Examination: Biology: 1998, © New Zealand Qualifications Authority 1998, Wellington, NZ. Extract only. Table Flowering in short-day and long-day plants; sources not given, from University Entrance, Bursaries and Scholarships Examination: Biology: 1998, © New Zealand Qualifications Authority 1998, Wellington, NZ. Extract only. Illustration All illustrations and diagrams copyright © Te Aho o Te Kura Pounamu, Wellington, NZ. BY3031A Photos Muehlenbeckia complexa climbing a tree; Muehlenbeckia complexa hugging the ground, © Claire Neiman, Wellington, NZ: Te Aho o Te Kura Pounamu. Used by permission. Wire vine (Muehlenbeckia complexa), in flower, Tiritiri Matangi Island, Wikimedia Commons, http://en.wikipedia.org/wiki/ File: Muehlenbeckia_complexia_in_flower_T2i_IMG_104_1452.jpg, accessed October 2012. Public domain. Cross section of a fig, by Rainer Zenz, Wikimedia Commons, http://en.wikipedia.org/wiki/File:Feige-Schnitt.jpg, accessed September 2012. Public domain. © te ah o o t e k ur a p o un a m u BY3031 65 pta tio ic env n di a r na di u ro di io er ph ot op an pl us im ul st d p us rio oc h t y h o en og ex lants top e ta t t n e m n o r i env me day p pho lants m e biotic short - bi day p ons ha long- sm m or th che ni l a c i g ecolo res p s endogenou is tic ent op tr nas all ironm l m cy n a on e ab sc is si on abiot r do y h t a elop n ci rc a au xi n ada op tr iation) c o s s a c i t s i utual m ( c i t o i b m sy per iod icit e c n a y r e l o t m is