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W
hile walking in the veld with Richard Cowling some time ago he asked me why Ithink some flowers close at night. My answer at the time was, ‘to protect their pol-len’, based on the simple fact that the pollen of most plant species loses its viability when it gets wet. So closing your petals to protect your pollen from night dew - andon cool rainy days- makes good sense as a lack of viable pollen will nullify the entirereason for flowering.But there is more to the story than my initial simple answer. Ever since Richardasked me that question I have been looking more carefully at flower behaviour andI would like you to join me to do just that. Flower behaviour? Yes, just like animals,plants also have a distinctive behaviour. They may not behave as individuals, butdifferent species certainly behave differently in terms of their flowers. The flowersof some species remain open day and night, in others they only open during the warmer parts of a day, whereas some species only open at night. You all knew that,but think carefully about these three behavioral patterns again. Why do plantsbehave differently?So, why do some flowers close in the evening and when it is cool during the day?A point clearly made in some recent articles in
Veld & Flora 
 
indicated that certainplants only open their flowers when it is warm. The implication here is that plantsget a signal as soon as the temperature drops in the evening, or when it remainscool during the day, that moisture is on its way. As soon as the temperature drops
while others stay open until all hours,come rain or shine?
Why do some flowers close at night
...
Watsonias often flower with a spectacular show after fire, but these plants have a problem. Few
Watsonia 
species have any particular flower design or flowering pattern strategy to protect their pol-len against moisture - the pollen killer. Like
Watsonia knysnana 
shown here, they seem to have optedfor an
en masse
flowering display strategy, a sort of 'Hitler model', hoping for the best even if theyknow that few flowers may survive to produce seed after inclement weather.
Photo: Jan Vlok.
 Jan Vlok 
looks at some intriguing examples of flower behaviour.
Plant behaviour
June 2005
Veld
&
Flora
76
ABOVE:
Whiteheadia bifolia 
employs a rather clever pagoda-like design to channel any rain-water away from the open flowers. A similar design also occurs in some
Eucomis 
species.Photo: Jan Vlok.
 
in the evening the relative humidity of the air increases, or rain may be immi-nent if it remains cool during the day.So cool air indicates to plants that thereis a risk that their pollen may becomemoist and it is best to close shop toprevent damage to goods. A secondreason for closing shop is because nocustomers are on the street. The activ-ity of most diurnal insects decreasesat low temperatures, which for flowersmeans no clients - so just stay in bed allday and don’t waste your energy. Justthe opposite happens with nocturnalinsects though. If, as a plant, you optfor nightlife clients, the evening signalto open your shop comes at a risk. The risk is losing the male function(production and storage of pollen inthe stamen) of your flower: little pointin having many pretty girls around inthe evening without having some viableboys around. Several of our plants usethis seemingly risky nightlife option,like the aandblommetjies (
Hesperantha 
 species). It is quite obvious why theseplants open their flowers in the evening:to attract specific pollinators, but notvery clear how they protect their pollenassets in the process, when other flow-ers tuck them up safe and dry. There isclearly more to the story than just theobvious.
Wash out
 The first time I noted how moisturecan affect pollination in plants was in aspectacular stand of 
Watsonia fourcadei 
 flowers on the slopes of the Outeniquamountains. I tried to collect pollen fromthese plants a day after rain, but couldnot find any, even though the flowers were perfectly intact. The rain washedaway all the pollen that was releasedbefore the rain and none of the anthersof the other open flowers were willing torelease their pollen: they were waitingfor warmer weather. No pollen was thusavailable in this population for two tothree days. Being puzzled by this I thenmarked the sections on the inflores-cences that had open flowers with bitsof wool and returned a few weeks later.Almost none of the flowers that wereopen at the time of the rain had devel-oped capsules with viable seed, simplybecause there was no viable pollen tobe pollinated. I have noted this subse-quently many times in
Watsonia 
and
Aloe 
inflorescences. Once the plantsare in seed one can even often tell whena cold front passed through the area,as a series of undeveloped capsulesmark those rainy days. I suspect thatthe same happens in
Protea 
flowers.I recently saw another interest-ing example in my garden when Iopened my sprinkler system to watera lovely bed of flowering
Nerine fili-  folia 
. Not a single one of them pro-duced seed afterwards, as I unthink-ingly washed all the viable pollen away.So we do have some species in whichthe flowers seem to offer little protec-tion to protect their pollen from cool, wet conditions. These species seemto be characterized by inflorescencesthat produce many flowers that opensequentially over a prolonged period.So, those flowers that are literallycaught with their pants down in wetconditions are simply compensated forby others that will open during betterconditions.
Staying dry
By now some of you will probablyalready mumble that this is just anoth-er ‘old-wives tale’ with no scientific evi-
TOP RIGHT: Safety in numbers.
Protea pudens 
is anexample of a plant that does not seem to employspecial measures to protect its pollen from gettingwet. The inflorescence, however, consists of manyflowers that are opened sequentially to ensure thatnew pollen is released after the rain has passed.BELOW RIGHT:
Gladiolus rogersii 
is a classic exam-ple of a ‘windsock-flower’, where a thin flexibleflowering stem enables the flower to turn its back on any moisture-laden wind. This flower designoften occurs in plants of high rainfall areas, whereit allows the flowers to stay open at night or evenwhen it rains because the pollen is very well pro- tected against moisture.
Photos: Jan Vlok.
LEFT:
Aloe ferox,
 
like most other aloes, never closes its flowers. None, or very few, of theseopen flowers will set seed if it rains. Aloes oftenflower in the dry season but even if it rains, theyjust rapidly open a whole set of new flowers after  the rain, and some will be pollinated. Once theplants are in seed one can even often tell that acold front passed through, as a series of undevel-oped capsules mark those rainy days.
Photo: Jan Vlok.
June 2005
Veld
&
Flora
77
LEFT: Most of the carrion flowers, like this
Piaranthus comptus,
stay open at night and evenwhen it rains. Like orchids, they have little toworry about as the viability of their pollen is safe:well protected in waterproof bags.
Photo: Jan Vlok.
 
dence. Well there is a way to test if flow-ers close when it cools down to protecttheir pollen from becoming wet. We cando this by comparing the flower behav-ior of species that have ‘waterproof’ pollen types with those that do not. The predicted outcome being that thosespecies that do protect their pollen in‘waterproof’ bags, or any other specificdesign of the anthers, would not closeat night. Perhaps the best examples of plants that retain their pollen in water-proof bags are orchids and carrion-flowers (stapeliads). Thus far I havenot seen any of our local orchids orstapeliads close their flowers at night orduring inclement weather. Once open,they remain open even though manyof them have only diurnal pollinators.Another group of interesting plantsthat we can use to test the idea are the‘buzz-pollination’ plants. This remark-able group of plants has anthers thatonly release their pollen when a pollina-tor ‘buzzes’ at the right frequency closeto them. Local examples include speciessuch as lady’s hand (
Cyanella 
species)and bitter apples (
Solanum 
species). Asin the case of our orchids and carrionflowers I have not yet seen any of thesebuzz-pollinated plants close their flow-ers at night.
Erica 
species are not buzz-pollinatedplants but they often only release theirpollen through small apical pores andas far as I know none of them closetheir flowers at night or during wet weather. So there are all sorts of antherdesigns that plants have discovered toovercome the wet pollen problem, butthere is more to the story. I believesome plants also have specific flowerdesigns to overcome this problem.An interesting example of special flow-er design occurs amongst the vygies (iceplants). Flowers of the genera
Erepsia 
 and
Smicrostigma 
are unusual amongstthe vygies as they do not close theirflowers at night. Why not? They arethe only vygies that hide their anthersunderneath a tightly closed cone of staminodes (filaments without anthers)that is watertight. To pollinate thesevygie flowers, beetles actually have to
ABOVE:
Brunsvigia josephiniae 
is one of many plants that onlyflowers in the dry season, usually in arid areas. Their flowersremain open in the evening, but as it is usually dry, there islittle risk of their pollen getting wet.
Photo: Jan Vlok.
FAR LEFT:If you ever wondered why orchid flowers do not close atnight, I would say that it is because their pollen is well pro- tected in little waterproof bags. Those flowering during therainy season, like the ewwa-trewwa
Satyrium coriifolium,
 seem to have taken extra protective measures by having an‘anther-umbrella’ flower design as well.LEFT: An interesting variation on the normal theme of flower display occurs amongst legumes such as
Aspalathus shawii.
 None of these close at night or during rain, simply because their pollen is well protected in the boat-shaped keel at thebase of the flower.
Photo: Jan Vlok.
June 2005
Veld
&
Flora
78
LEFT: What’s the buzz?
Cyanella orchidi- formis 
is one of the remarkable ‘buzz-pol-linated’ plants that only releases its pollen through a narrow pore when an insectbuzzes at the correct frequency close by.The pollen is naturally protected againstmoisture and the flowers remain open incool conditions.FAR LEFT:
Erepsia pentagona 
is an unu-sual vygie that remains open at night andeven during rain. The tight cone of stami-nodes
 
(filaments without anthers) in thecentre of the flower covers the anthers andensures that no moisture can reach thepollen.
Photos: Jan Vlok.
Some plants may even dictateto their pollinators how theyshould behave
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