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Fluid Physiology

Fluid Physiology:

A Handbook for Anaesthesia

and Critical Care Practice

By

Thomas Woodcock
Fluid Physiology: A Handbook for Anaesthesia and Critical Care Practice

By Thomas Woodcock

This book first published 2019

Cambridge Scholars Publishing

Lady Stephenson Library, Newcastle upon Tyne, NE6 2PA, UK

British Library Cataloguing in Publication Data

A catalogue record for this book is available from the British Library

Copyright © 2019 by Thomas Woodcock

All rights for this book reserved. No part of this book may be reproduced,
stored in a retrieval system, or transmitted, in any form or by any means,
electronic, mechanical, photocopying, recording or otherwise, without
the prior permission of the copyright owner.

ISBN (10): 1-5275-4031-6

ISBN (13): 978-1-5275-4031-6
 



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ŚĂƉƚĞƌ^ƵŵŵĂƌLJ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϰϯ
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DŝƚŽĐŚŽŶĚƌŝĂůŚLJĚƌŽŐĞŶŝŽŶŐƌĂĚŝĞŶƚ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϰϲ
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sŽůƵŵĞĂŶĚƚŚŝĐŬŶĞƐƐŽĨƚŚĞĞŶĚŽƚŚĞůŝĂůƐƵƌĨĂĐĞůĂLJĞƌ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϴϳ
dŚĞĞŶĚŽƚŚĞůŝĂůƐƵƌĨĂĐĞůĂLJĞƌ͖ŝŶĨůĂƚŝŽŶĂŶĚĚŝƐŝŶƚĞŐƌĂƚŝŽŶ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϴϵ
&ƵŶĐƚŝŽŶƐŽĨƚŚĞĞŶĚŽƚŚĞůŝĂůŐůLJĐŽĐĂůLJdž͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϵϭ
sĂƐĐƵůĂƌĞŶĚŽƚŚĞůŝĂůĐĞůůƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϵϱ
/ŶƚĞƌĞŶĚŽƚŚĞůŝĂůũƵŶĐƚŝŽŶƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϵϱ
ƋƵĂƉŽƌŝŶĐŚĂŶŶĞůƐĂƌĞƉƌĞƐĞŶƚŝŶĞŶĚŽƚŚĞůŝĂůĐĞůůƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϵϳ
ƋƵĂƉŽƌŝŶĐŚĂŶŶĞůƐŽĨĂƐƚƌŽĐLJƚĞĨŽŽƚƉƌŽĐĞƐƐĞƐĂƌĞŝŵƉŽƌƚĂŶƚƚŽ
ďůŽŽĚͲďƌĂŝŶƉĞƌŵĞĂďŝůŝƚLJďĂƌƌŝĞƌ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϵϴ
ƋƵĂƉŽƌŝŶϮŝƐĞdžƉƌĞƐƐĞĚŝŶƚŚĞƌĞŶĂůĐŽůůĞĐƚŝŶŐĚƵĐƚ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϵϴ
ƋƵĂƉŽƌŝŶƐŝŶƚŚĞůƵŶŐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϵϴ
ĂƉŝůůĂƌLJĐůĂƐƐĞƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϵϴ
ŝƌĐƵůĂƚŝŶŐĞŶĚŽƚŚĞůŝĂůĐĞůůƐĂŶĚĞŶĚŽƚŚĞůŝĂůͲĚĞƌŝǀĞĚŵŝĐƌŽƉĂƌƚŝĐůĞƐϭϬϱ
dŚĞƐƵďĞŶĚŽƚŚĞůŝĂůďĂƐĞŵĞŶƚŵĞŵďƌĂŶĞĂŶĚĞdžƚƌĂĐĞůůƵůĂƌŵĂƚƌŝdž ϭϬϱ
ZĞĨĞƌĞŶĐĞƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϬϲ
 
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ŚĂƉƚĞƌϱ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϭϮ
/ŶƚĞƌƐƚŝƚŝƵŵĂŶĚ>LJŵƉŚĂƚŝĐ^LJƐƚĞŵƐ
ŚĂƉƚĞƌƐƵŵŵĂƌLJ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϭϮ
/ŶƚĞƌƐƚŝƚŝƵŵ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϭϮ
DŽůĞĐƵůĂƌƐƚƌƵĐƚƵƌĞŽĨƚŚĞƚƌŝƉŚĂƐŝĐŝŶƚĞƌƐƚŝƚŝƵŵ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϭϯ
,LJĂůƵƌŽŶĂŶ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϭϰ
dŚĞƉĞƌŝǀĂƐĐƵůĂƌŵĂƚƌŝdž͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϭϱ
/ŶƚĞƌƐƚŝƚŝĂůǁĂƚĞƌ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϭϲ
'ĞůƐǁĞůůŝŶŐƉƌĞƐƐƵƌĞ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϭϳ
ŽůůŽŝĚŽƐŵŽƚŝĐƉƌĞƐƐƵƌĞŽĨƚŚĞŝŶƚĞƌƐƚŝƚŝƵŵ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϭϳ
ůďƵŵŝŶĞdžĐůƵƐŝŽŶĂŶĚĂƋƵĞŽƵƐƉŚĂƐĞǀŝƐĐŽƐŝƚLJ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϭϳ
/ŶƚĞƌƐƚŝƚŝĂůĨůƵŝĚƉƌĞƐƐƵƌĞ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϭϴ
ŶĂƚŽŵŝĐĨĞĂƚƵƌĞƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϭϴ
>LJŵƉŚĂƚŝĐǀĂƐĐƵůĂƌƐLJƐƚĞŵ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϭϵ
/ŶƚĞƌƐƚŝƚŝĂůĨůƵŝĚĚLJŶĂŵŝĐƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϮϬ
>LJŵƉŚĂƚŝĐƐĂŶĚƚŚĞŝŶƚĞƌƐƚŝƚŝĂůƐƚŽƌĂŐĞŽĨƐŽĚŝƵŵ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϮϮ
/ŶƚĞƌƐƚŝƚŝĂůĨůƵŝĚŝŶĐƌŝƚŝĐĂůŝůůŶĞƐƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϮϰ
WĞƌŝƚŽŶĞĂůĚŝĂůLJƐŝƐ͕ƉĞƌŝƚŽŶĞĂůƌĞƐƵƐĐŝƚĂƚŝŽŶ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϮϲ
ZĞĨĞƌĞŶĐĞƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϮϲ

ŚĂƉƚĞƌϲ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϯϬ
^LJƐƚĞŵŝĐĐŝƌĐƵůĂƚŝŽŶ͗dŚĞƉĞƌŝƉŚĞƌĂůǀĂƐĐƵůĂƌůŽŽƉƐ
ŚĂƉƚĞƌƐƵŵŵĂƌLJ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϯϬ
^LJƐƚĞŵŝĐĐŝƌĐƵůĂƚŝŽŶ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϯϬ
ĞƌĞďƌĂůĐŝƌĐƵůĂƚŝŽŶ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϯϬ
ŽƌŽŶĂƌLJĐŝƌĐƵůĂƚŝŽŶ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϯϱ
,ĞƉĂƚŝĐĐŝƌĐƵůĂƚŝŽŶƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϯϲ
/ŶƚĞƐƚŝŶĂůŵƵĐŽƐĂůĐŝƌĐƵůĂƚŝŽŶ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϯϳ
/ŶƚĞƌƐƚŝƚŝĂůĨůƵŝĚĂŶĚ>LJŵƉŚŝƌĐƵůĂƚŝŽŶŝŶƚŚĞ>ŝǀĞƌĂŶĚŝŶƚĞƐƚŝŶĂů
ŵƵĐŽƐĂ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϯϴ
^ƉůĞŶŝĐĐŝƌĐƵůĂƚŝŽŶ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϯϵ
ŽŶĞŵĂƌƌŽǁĐŝƌĐƵůĂƚŝŽŶ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϰϬ
ZĞŶĂůĐŝƌĐƵůĂƚŝŽŶ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϰϬ
^ĞĐƌĞƚŽƌLJŐůĂŶĚƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϰϯ
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ZĞĨĞƌĞŶĐĞƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϰϰ
 
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WůĞƵƌĂůĨůƵŝĚ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϱϱ
ZĞĨĞƌĞŶĐĞƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϱϳ

ŚĂƉƚĞƌϴ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϱϵ
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/ŶƚƌŽĚƵĐƚŝŽŶ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϱϵ
ĞĨŝŶŝŶŐĂŶĚŵĞĂƐƵƌŝŶŐƉůĂƐŵĂǀŽůƵŵĞ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϲϬ
sŽůƵŵĞ^ĞŶƐŽƌƐĂŶĚŶĞƵƌŽŚƵŵŽƌĂůƌĞƐƉŽŶƐĞ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϲϯ
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ƌĞǀŝƐĞĚdǁŝŐůĞLJͲ,ŝůůŵĂŶĚŝĂŐƌĂŵ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϳϬ
dŚĞƌĂƉĞƵƚŝĐŽƉƚŝŽŶƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϳϮ
ZĞĨĞƌĞŶĐĞƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϳϰ

ŚĂƉƚĞƌϵ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϳϳ
ůďƵŵŝŶĂŶĚKƚŚĞƌŝƌĐƵůĂƚŝŶŐWƌŽƚĞŝŶƐ
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ůďƵŵŝŶ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϳϴ
ŝƌĐƵůĂƚŝŽŶŽĨĂůďƵŵŝŶ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϳϴ
ůŝŶŝĐĂůŵĞĂƐƵƌĞŵĞŶƚŽĨĂůďƵŵŝŶ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϴϬ
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ůďƵŵŝŶĂƐĂďŝŶĚŝŶŐƉƌŽƚĞŝŶ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϴϯ
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ůďƵŵŝŶĂŶĚĐĂƉŝůůĂƌLJƉĞƌŵĞĂďŝůŝƚLJ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϴϯ
ůďƵŵŝŶŝŶŵĂůŶƵƚƌŝƚŝŽŶ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϴϱ
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ůďƵŵŝŶŝŶƉƌĞŐŶĂŶĐLJ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϴϲ
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ĐƵƚĞWŚĂƐĞWƌŽƚĞŝŶƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϴϴ
ZĞĨĞƌĞŶĐĞƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϴϵ

ŚĂƉƚĞƌϭϬ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϵϯ
ŚĂĞŵŽĚLJŶĂŵŝĐƉĂƌĂĚŝŐŵ
ŚĂƉƚĞƌƐƵŵŵĂƌLJ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϵϯ
,ŝƐƚŽƌŝĐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϵϯ
dŚĞĐƵƌƌĞŶƚůLJͲƉƌĞǀĂŝůŝŶŐƉĂƌĂĚŝŐŵ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϵϲ
ĂƌĚŝŽŵLJƚŚŽůŽŐLJ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϵϳ
DĞĂŶĐŝƌĐƵůĂƚŽƌLJĨŝůůŝŶŐƉƌĞƐƐƵƌĞ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϵϴ
sĞŶŽƵƐĞdžĐĞƐƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϭϵϵ
DLJŽĐĂƌĚŝĂůĞŶĞƌŐLJĞdžĐĞƐƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϮϬϮ
ĨĨĞĐƚŝǀĞǀĞƌƐƵƐŝŶĞĨĨĞĐƚŝǀĞďůŽŽĚǀŽůƵŵĞƐ͍͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϮϬϱ
ĞƐĐƌŝďŝŶŐƉƌĞƐƐƵƌĞͲǀŽůƵŵĞĐŚĂƌĂĐƚĞƌŝƐƚŝĐƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϮϬϳ
ƌƚĞƌŝĂůƉƌĞƐƐƵƌĞͲǀŽůƵŵĞĐŚĂƌĂĐƚĞƌŝƐƚŝĐƐŝŶƉƌĂĐƚŝĐĞ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϮϬϴ
^LJƐƚĞŵŝĐǀĂƐĐƵůĂƌǁĂƚĞƌĨĂůů͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϮϭϬ
WƌĞƐƐƵƌĞͲǀŽůƵŵĞĂŶĚĨůŽǁĐŚĂƌĂĐƚĞƌŝƐƚŝĐƐŽĨƚŚĞǀĂƐĐƵůĂƌůŽŽƉ͘͘͘͘͘͘͘Ϯϭϭ
,ĂĞŵŽĚLJŶĂŵŝĐĐŽŵƉƵƚĂƚŝŽŶŵŽĚĞůƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘Ϯϭϲ
^ŚŽĐŬ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘Ϯϭϳ
ZĞĨƌĂĐƚŽƌLJƐŚŽĐŬ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘Ϯϭϳ
^ŚŽĐŬŝŶŚŝůĚƌĞŶ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘Ϯϭϴ
ƐƐĞƐƐŝŶŐƐŚŽĐŬĂŶĚƚŚĞƌĞƐƉŽŶƐĞƚŽƚƌĞĂƚŵĞŶƚ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘Ϯϭϴ
,ĞĂƌƚĨĂŝůƵƌĞ͖ĂŶĂƉƉƌŽƉƌŝĂƚĞŶĂŵĞĨŽƌǀĞŶŽƵƐĐŽŶŐĞƐƚŝŽŶ͍͘͘͘͘͘͘͘͘͘͘͘͘Ϯϭϵ
ZĞͲĞǀĂůƵĂƚŝŶŐĐĞŶƚƌĂůǀĞŶŽƵƐƉƌĞƐƐƵƌĞŵŽŶŝƚŽƌŝŶŐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϮϮϭ
ďĞƚƚĞƌŚĂĞŵŽĚLJŶĂŵŝĐƉĂƌĂĚŝŐŵ͍͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϮϮϭ
ZĞĨĞƌĞŶĐĞƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϮϮϯ

ŚĂƉƚĞƌϭϭ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϮϮϵ
ŶŝŵƉƌŽǀĞĚƉĂƌĂĚŝŐŵŽĨ&ůƵŝĚWŚLJƐŝŽůŽŐLJĂŶĚdŚĞƌĂƉLJ
ŚĂƉƚĞƌƐƵŵŵĂƌLJ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϮϮϵ
ǀĂůŝĚĐůŝŶŝĐĂůƉĂƌĂĚŝŐŵŽĨďŽĚLJǁĂƚĞƌĚŝƐƉŽƐŝƚŝŽŶĂŶĚƌĞƐƉŽŶƐĞƚŽ
ŝŶƚƌĂǀĞŶŽƵƐŝŶĨƵƐŝŽŶƐŵƵƐƚĞdžƉůĂŝŶŽďƐĞƌǀĞĚĨĂĐƚƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϮϮϵ
DŽŽƌĞ͛ƐsŽůƵŵĞ<ŝŶĞƚŝĐƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϮϯϬ
dŚĞƌĂƚŝŽŽĨƉůĂƐŵĂǀŽůƵŵĞƚŽŝŶƚĞƌƐƚŝƚŝĂůĨůƵŝĚǀŽůƵŵĞ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘Ϯϯϭ
dŚĞƌĂƚĞŽĨƉůĂƐŵĂǀŽůƵŵĞƌĞĨŝůůĂĨƚĞƌďůŽŽĚůŽƐƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘Ϯϯϭ
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sŽůƵŵĞŬŝŶĞƚŝĐƐŽĨĂŶŝŶĨƵƐĞĚŝƐŽƚŽŶŝĐƐĂůƚƐŽůƵƚŝŽŶ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϮϯϮ
ƌLJƐƚĂůůŽŝĚŝŶĨƵƐŝŽŶĞŶŚĂŶĐĞƐƚŚĞŵŽǀĞŵĞŶƚŽĨŝŶƚĞƌƐƚŝƚŝĂůƉƌŽƚĞŝŶƐ
ƚŽƚŚĞƉůĂƐŵĂ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϮϯϮ
DŽŽƌĞ͛ƐĨŽƵƌƌĂƚĞĨĂĐƚŽƌƐƚŽĐŽŶƐŝĚĞƌ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘Ϯϯϯ
dŚĞĚĞǀĞůŽƉŵĞŶƚŽĨǀŽůƵŵĞŬŝŶĞƚŝĐƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘Ϯϯϯ
,ĂŚŶ͛ƐsŽůƵŵĞ<ŝŶĞƚŝĐƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘Ϯϯϰ
,ĂŚŶ͛ƐƌĂƚĞĨĂĐƚŽƌƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘Ϯϯϱ
ƉƉůŝĐĂƚŝŽŶƐŽĨ,ĂŚŶ͛ƐǀŽůƵŵĞŬŝŶĞƚŝĐƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘Ϯϯϲ
WŚĂƌŵĂĐŽĚLJŶĂŵŝĐƐŽĨĂĨůƵŝĚĐŚĂůůĞŶŐĞ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘Ϯϯϴ
&ƵƚƵƌĞǀŽůƵŵĞŬŝŶĞƚŝĐƌĞƐĞĂƌĐŚ͍͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘Ϯϯϵ
dŚĞ'ůLJĐŽĐĂůLJdžDŽĚĞůWĂƌĂĚŝŐŵƚŚĞŶĂŶĚŶŽǁ͗dĞƌŵŝŶŽůŽŐLJ͘͘͘͘͘͘͘͘͘͘Ϯϯϵ
dŚĞŝŶƚƌĂǀĂƐĐƵůĂƌƐƉĂĐĞ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϮϰϬ
dŚĞŚĞƚĞƌŽŐĞŶĞŝƚLJŽĨĐĂƉŝůůĂƌŝĞƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘Ϯϰϭ
dŚĞ^ƚĂƌůŝŶŐĨŽƌĐĞƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϮϰϮ
dŚĞ:ĐƵƌǀĞ͖ĂŚŽĐŬĞLJƐƚŝĐŬ͍͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϮϰϮ
dŚĞĞĨĨĞĐƚƐŽĨĐŽůůŽŝĚĂŶĚĐƌLJƐƚĂůůŽŝĚŝŶĨƵƐŝŽŶƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘Ϯϰϯ
dŚĞŝŶƚĞƌƐƚŝƚŝƵŵĂŶĚůLJŵƉŚĂƚŝĐƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘Ϯϰϰ
ƚǁŽͲĐŽŵƉĂƌƚŵĞŶƚŬŝŶĞƚŝĐŵŽĚĞů͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘Ϯϰϱ
ƌĞǀŝƐĞĚdǁŝŐůĞLJͲ,ŝůůŵĂŶĚŝĂŐƌĂŵ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘Ϯϰϴ
ŶŝĚĞĂůĞdžƚƌĂĐĞůůƵůĂƌĨůƵŝĚƐƵďƐƚŝƚƵƚĞ͍͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϮϱϬ
ŽŶƚĞdžƚƐĞŶƐŝƚŝǀŝƚLJĂŶĚsŽůƵŵĞĞƋƵŝǀĂůĞŶĐĞ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘Ϯϱϭ
hŶĚĞƌƐƚĂŶĚŝŶŐůĞĂŬLJĐĂƉŝůůĂƌŝĞƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘Ϯϱϰ
ŐůLJĐŽĐĂůLJdžŵŽĚĞůƉĂƌĂĚŝŐŵŽĨŚĂĞŵŽĚLJŶĂŵŝĐĐŽŶƚƌŽů͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘Ϯϱϳ
ŝƐĞƋƵŝůŝďƌŝƵŵĞǀĞŶƚƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘Ϯϱϴ
ŶĂůƚĞƌŶĂƚŝǀĞůŽŽŬĂƚƐLJƐƚĞŵŝĐŝŶĨůĂŵŵĂƚŽƌLJŚĂĞŵŽĚLJŶĂŵŝĐƐ͘͘͘͘͘͘͘Ϯϱϵ
ŽŶĐůƵĚŝŶŐƌĞŵĂƌŬƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘ϮϲϬ
ZĞĨĞƌĞŶĐĞƐ͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘͘Ϯϲϭ





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‹‡ ƒ›ǡ  ™ƒ• ‹–”‘†— ‡† –‘ ”Ǥ Š‘ƒ• ‘‘† ‘ ǡ Dz‘dzǡ „›
”‡ƒ†‹‰ Š‹• ’—„Ž‹ ƒ–‹‘ ‘ˆ –Š‡ ‡˜‹•‡† –ƒ”Ž‹‰ ‡“—ƒ–‹‘ ƒ† –Š‡
‰Ž› ‘ ƒŽ›š‘†‡Ž‘ˆ–”ƒ•˜ƒ• —Žƒ”ˆŽ—‹†‡š Šƒ‰‡ǤŠ‘”–Ž›–Š‡”‡ƒˆ–‡” 
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Ž‡ –—”‡ ƒ– Š‡ Š›•‹‘Ž‘‰‹ ƒŽ ‘ ‹‡–›ǡ ‘†‘ʹ ™Š‡”‡‹ Š‡ ™‹––‹Ž› ƒ†
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ˆŽ—‹††›ƒ‹ •ǤŠ‘”–Ž›–Š‡”‡ƒˆ–‡” Šƒ†–Š‡‘’’‘”–—‹–›–‘‹˜‹–‡‘
–‘ –Š‡ ʹ͵”† –‡”ƒ–‹‘ƒŽ ‡–‡”‹ƒ”› ‡”‰‡ › ƒ† ”‹–‹ ƒŽ ƒ”‡
›’‘•‹—ȋ ȌŠ‡Ž†ƒ––Š‡ ƒ›Ž‘”†’”›Žƒ†‡•‘”–‹ƒ•Š˜‹ŽŽ‡
ǡ ‡’–‡„‡” ʹͲͳ͹Ǥ Š‹• ™ƒ• ™‹–Š‘—– “—‡•–‹‘ ‘‡ ‘ˆ –Š‡ ‘•–
‡Ž‹‰Š–‡‹‰ ƒ† ƒŽŽǦ–‘‘ „”‹‡ˆ ‡ ‘—–‡”•  Šƒ˜‡ ‡Œ‘›‡† ™‹–Š ‘
‘–Š‡” –Šƒ ‘—” ‘‰‘‹‰ ‡ƒ‹Ž ‡š Šƒ‰‡• ’‘†‡”‹‰ ˜ƒ• —Žƒ”
’Š›•‹‘Ž‘‰›ǡ ˆŽ—‹† ’Š›•‹‘Ž‘‰›ǡ ˆŽ—‹† „ƒŽƒ ‡ ƒ† ˆŽ—‹† –Š‡”ƒ’›Ǥ  ›
‘’‹‹‘ ‘ ‹• ‘‡ ‘ˆ ƒ Šƒ†ˆ—Ž ‘ˆ Ž‹‹ ƒŽ • ‹‡–‹•–• –Šƒ– ‹• ƒ„Ž‡ –‘
‘ ‡’–—ƒŽ‹œ‡ ƒ† –”ƒ•Žƒ–‡ ‘’Ž‡šǡ ƒ† ƒ– –‹‡•ǡ ˜ƒ‰—‡ ƒ†
ƒ„‹‰—‘—• ’Š›•‹‘Ž‘‰‹  ’”‘ ‡••‡• ‹–‘ –”ƒ•’ƒ”‡– • ‡ƒ”‹‘• –Šƒ–
‡š’Žƒ‹ –Š‡‹” Ž‹‹ ƒŽ ”‡Ž‡˜ƒ ‡ ƒ† –Š‡”ƒ’‡—–‹  ‹’Ž‹ ƒ–‹‘•Ǥ ‘ǯ•
™”‹–‹‰•ǡŽ‡ –—”‡•ǡƒ†™‡„•‹–‡’‘•–•Šƒ˜‡‡Ž‹‰Š–‡‡†ƒ†‘†‡”‹œ‡†
‘—” —†‡”•–ƒ†‹‰ ‘ˆ ˜ƒ• —Žƒ” ƒ† ˆŽ—‹† ’Š›•‹‘Ž‘‰› ƒ† Š‹‰ŠŽ‹‰Š–
— ‡”–ƒ‹–‹‡• –Šƒ– ƒ ‡™ ‰‡‡”ƒ–‹‘ ‘ˆ ˆŽ—‹† –Š‡”ƒ’› ‡–Š—•‹ƒ•–• ƒ
‹˜‡•–‹‰ƒ–‡Ǥ‘ǯ•–‡ƒ Š‹‰ƒ† Ž‹‹ ƒŽ‡š’‡”‹‡ ‡•‹ ‘Œ— –‹‘™‹–Š
Š‹• ‡‡ ‹–‡”‡•– ‹ ˆŽ—‹† ˆŽ—šǡ ˆŽ—‹† ˆ‹Ž–”ƒ–‹‘ǡ ƒ† ˆŽ—‹† „ƒŽƒ ‡
‡ ‘’ƒ••‘†‡”†ƒ›‡˜‹†‡ ‡Ǧ„ƒ•‡††‹• ‘˜‡”‹‡•‡˜‡”ƒ†˜ƒ ‹‰Š‹•
’”‘‘–‹‘‘ˆ–Š‡Dz”‡˜‹•‡†–ƒ”Ž‹‰’”‹ ‹’Ž‡dzǤŠ‹•”‡˜‹•‹‘ ŠƒŽŽ‡‰‡•
Š‹•–‘”‹ ƒŽ ‹–‡”’”‡–ƒ–‹‘• ƒ† ‡ ‘—”ƒ‰‡• –Š‡ †‡˜‡Ž‘’‡– ‘ˆ ƒ ‡™
’ƒ”ƒ†‹‰ ˆ‘” ˆŽ—‹† •‡Ž‡ –‹‘ ƒ† ƒ†‹‹•–”ƒ–‹‘ǡ ‡•’‡ ‹ƒŽŽ› ”‡‰ƒ”†‹‰
–Š‡ƒ†‹‹•–”ƒ–‹‘‘ˆ„‹‘’Š›•‹ ƒŽ–Š‡”ƒ’‹‡•Ǥ‘•–‘–ƒ„Ž›ǡƒ† ‘–”ƒ”›
–‘ƒ›–‡š–„‘‘•ǡ–Š‹•’ƒ”ƒ†‹‰’”‡†‹ –•–Šƒ–ƒ–•—„‘”ƒŽƒ”–‡”‹ƒŽ
ƒ† ƒ’‹ŽŽƒ”›’”‡••—”‡–Š‡‹ˆ—•‹‘‘ˆƒ‹•‘–‘‹  ”›•–ƒŽŽ‘‹†•‘Ž—–‹‘
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ˆŽ—‹† ˆŽ—š ”‡ƒ‹• Ž‘•‡ –‘ œ‡”‘ ƒ†ǡ ‘–”ƒ”› –‘ ‘•– –‡š–„‘‘•ǡ –Š‡
ƒ†‹‹•–”ƒ–‹‘‘ˆƒ ‘ŽŽ‘‹†•‘Ž—–‹‘’”‘˜‹†‡•Ž‹––Ž‡–‘‘•‹‰‹ˆ‹ ƒ–Ž‘‰Ǧ

ͳŠ––’•ǣȀȀˆŽ—‹†’Š›•‹‘Ž‘‰›Ǥ‘”‰Ȁ
ʹŠ––’•ǣȀȀ™™™Ǥ›‘—–—„‡Ǥ ‘Ȁ™ƒ– Šǫ˜α‘ ˜˜ –—ǡ
 Ž—‹†Š›•‹‘Ž‘‰› š‹‹‹

–‡”–Š‡”ƒ’‡—–‹ ƒ†˜ƒ–ƒ‰‡•Ǥ‡ƒ†‡”•™‹ŽŽ‡Œ‘›ƒ Ž‡ƒ”ǡ ‘ ‹•‡ǡƒ†

‘’”‡Š‡•‹„Ž‡†‡• ”‹’–‹‘‘ˆ„‘†›ˆŽ—‹† ‘’ƒ”–‡–•ǡ–Š‡‡›”‘Ž‡
‘ˆ –Š‡ ‡†‘–Š‡Ž‹ƒŽ •—”ˆƒ ‡ Žƒ›‡” ȋ‹Ǥ‡Ǥ ‰Ž› ‘ ƒŽ›šȌ ƒ† –Š‡ ˜‹–ƒŽ
‹’‘”–ƒ ‡ ‘ˆ ‹–‡”•–‹–‹ƒŽ ˆŽ—‹† ƒ† Ž›’Š ˆŽ‘™ ‹ †‡–‡”‹‹‰
–”ƒ•˜ƒ• —Žƒ” ˆŽ—‹† ˆŽ—š ‹ „‘–Š –Š‡ ’—Ž‘ƒ”› ƒ† •›•–‡‹ 
‹” —Žƒ–‹‘•ǤŠ‡›™‹ŽŽƒŽ•‘‡Œ‘›Š‹•†‹• —••‹‘‘ˆ™Š›˜‡‘—•‡š ‡••
ƒ† ‘– ˜‡‘—• ”‡–—” ‹• –Š‡ ’”‹ƒ”› †‡–‡”‹ƒ– ‘ˆ ƒ”†‹ƒ  ‘—–’—–Ǥ
Š‡•‡†‹• —••‹‘•ƒ’’Ž›–‘ƒŽŽƒƒŽ•ƒ†™‹ŽŽ ƒ’–—”‡–Š‡‹–‡”‡•–‘ˆ
ƒ›‘‡ ‹–‡”‡•–‡† ‹ ˆŽ—‹† „ƒŽƒ ‡ ƒ† ˆŽ—‹† –Š‡”ƒ’›Ǥ  ‘ǯ• ™”‹–‹‰
•–›Ž‡ ‘„‹‡†™‹–ŠŠ‹•Ž— ‹†‡˜‹†‡ ‡Ǧ„ƒ•‡††‹• —••‹‘•‡ ‘—”ƒ‰‡•
”‡ƒ†‡”• –‘ ”‡–Š‹ ƒ› Š‹•–‘”‹  ˆŽ—‹† ’Š›•‹‘Ž‘‰› ’”‡ ‡’–• ƒ† –Š‡‹”
‹’Ž‹ ƒ–‹‘• ˆ‘” ˆŽ—‹† –Š‡”ƒ’›Ǥ ’‘”–ƒ–Ž› –Š‹• –‡š– ’”‘˜‹†‡• ƒ ‘”‡
’Š›•‹‘Ž‘‰‹  „ƒ•‡† —†‡”•–ƒ†‹‰ ‘ˆ „‘†› ˆŽ—‹† ‹‡–‹ • ƒ†
–”ƒ•˜ƒ• —Žƒ” ˆŽ—‹† ˆŽ—š –Šƒ– ‘ˆ‘”• –‘ Ž‹‹ ƒŽ ‘„•‡”˜ƒ–‹‘• ƒ†
†‹”‡ –• ”‡ƒ†‡”• –‘™ƒ”†• ‹’”‘˜‡†–Š‡”ƒ’‡—–‹ ƒ’’”‘ƒ Š‡• ‘ˆ ‘–‡š–
•’‡ ‹ˆ‹ ˆŽ—‹†„ƒŽƒ ‡†‹•‘”†‡”•ǤŠ‹•–‡š–‹•ƒ—•–”‡ƒ†ˆ‘”ƒŽŽ‡†‹ ƒŽ
’”‘ˆ‡••‹‘ƒŽ• ȋŠ—ƒ ƒ† ˜‡–‡”‹ƒ”›Ȍ ‹–‡”‡•–‡† ‹ ƒ†˜ƒ ‹‰ –Š‡‹”
‘™Ž‡†‰‡ƒ†—†‡”•–ƒ†‹‰‘ˆ–Š‡’Š›•‹‘Ž‘‰‹ ’”‘ ‡••‡•–Šƒ–†‹ –ƒ–‡
•ƒˆ‡ƒ†‡ˆˆ‡ –‹˜‡ˆŽ—‹†–Š‡”ƒ’›’”ƒ –‹ ‡•Ǥ 

 William MuirDVM, MS, PhD, DACVA, DACVECC, VCPC

͵͵ͺǤ͹–Š˜‡Ǥ
‘Ž—„—•ǡ Ͷ͵ʹͲͳ

”‘ˆ‡••‘” ‘ˆ Š›•‹‘Ž‘‰›ǡ ‘ŽŽ‡‰‡ ‘ˆ ‡–‡”‹ƒ”› ‡†‹ ‹‡ ƒ– ‹ ‘Ž
‡‘”‹ƒŽ‹˜‡”•‹–›‹ ƒ””‘‰ƒ–‡ǡƒ† ‹‡–‹ˆ‹ †˜‹•‘”ˆ‘”‡•–
Žƒ„‘”ƒ–‘”‹‡•ȋ’”‡ Ž‹‹ ƒŽ”‡•‡ƒ” Šˆƒ ‹Ž‹–›Ȍ‹‘Ž—„—• Ǥ

Previously;

”‘ˆ‡••‘”ƒ†‹”‡ –‘”‘ˆ‡•–Š‡•‹‘Ž‘‰›ƒ†ƒ‹ƒƒ‰‡‡–ƒ–Š‡
Š‹‘–ƒ–‡‹˜‡”•‹–›‘ŽŽ‡‰‡‘ˆ‡–‡”‹ƒ”›‡†‹ ‹‡ȋͳͻ͹ͲǦʹͲͲ͹Ȍ
‡•‡ƒ” Š ‹”‡ –‘” ƒ– ‡•‡ƒ” Š ‡†‹ ƒ–‹‘• ƒ† ‡•–‹‰ ‘•‘”–‹—
ȋʹͲͲ͹ǦͻȌ
Š‹‡ˆ‡†‹ ƒŽˆˆ‹ ‡”ǡ‹ƒŽ‡†‹ ƒŽ‡–‡”ǡȋʹͲͲͻǦͳʹȌ


  



„‡Ž‹‡˜‡–Šƒ–—†‡”•–ƒ†‹‰ˆŽ—‹†’Š›•‹‘Ž‘‰›‹• ”‹–‹ ƒŽ–‘–Š‡’”ƒ –‹ ‡
‘ˆ ‡†‹ ‹‡Ǥ ‘” ƒ› ›‡ƒ”•ǡ  ‰ƒ˜‡ Ž‡ –—”‡• ‘ ‹–”ƒ˜‡‘—• ˆŽ—‹†
–Š‡”ƒ’› –‘ ‹ƒŽ ‡ŽŽ‘™•Š‹’ ‡šƒ‹ƒ–‹‘ ƒ†‹†ƒ–‡• ƒ– –Š‡ ‘›ƒŽ
‘ŽŽ‡‰‡ ‘ˆ ƒ‡•–Š‡–‹•–• ‹ ‘†‘Ǥ Š‡ ’Š›•‹‘Ž‘‰‹ ƒŽ ’”‹ ‹’Ž‡• 
–ƒ—‰Š– ™‡”‡ ‡••‡–‹ƒŽŽ› –Š‘•‡ †‡• ”‹„‡† „› Ž‹•‘ ™‹‰Ž‡› ƒ† ‡
‹ŽŽƒ„ƒ ‹ͳͻͺ͸ȏͳȐǤ –™ƒ•–Š‡‹” Žƒ‹–Šƒ–‹ˆ›‘—™ƒ–‡†–‘‡š’ƒ†
–Š‡ ’Žƒ•ƒ ˜‘Ž—‡ǡ ›‘— •Š‘—Ž† ƒ†‹‹•–‡” ƒ ƒŽ„—‹ •‘Ž—–‹‘ ‘” ƒ
•›–Š‡–‹  ‘ŽŽ‘‹†•‘Ž—–‹‘Ǥ ˆ›‘—”ƒ‹™ƒ•–‘‡š’ƒ†–Š‡‡š–”ƒ ‡ŽŽ—Žƒ”
ˆŽ—‹† ˜‘Ž—‡ǡ ‰‹˜‡ ƒ ‹•‘–‘‹  •ƒŽ– •‘Ž—–‹‘ Ž‹‡ ‘”ƒŽ ƒŽ‹‡ ‘”
ƒ”–ƒǯ•‘Ž—–‹‘Ǥˆ”‡‡™ƒ–‡”ȋ‘”‘Ǧ‡Ž‡ –”‘Ž›–‡Ȍ•‘Ž—–‹‘ǡ•— Šƒ•
ͷΨ‡š–”‘•‡ǡ™‘—Ž††‹•–”‹„—–‡‹–•‡Žˆ–Š”‘—‰Š‘—––Š‡–‘–ƒŽ„‘†›™ƒ–‡”Ǥ
Š‡› ‘ Ž—†‡†–Šƒ––Š‹•”‡˜‡Žƒ–‹‘ƒ”‡†Dz–Š‡‡†‘ˆ–Š‡ ”›•–ƒŽŽ‘‹†
‡”ƒdz ƒ† –Šƒ– ‘”‡ ˆ‘ —• ‘ ’”‡•‡”˜‹‰ ’Žƒ•ƒ ‘ŽŽ‘‹† ‘•‘–‹ 
’”‡••—”‡ ‘—Ž† ”‡†— ‡ –Š‡ –‘–ƒŽ ˜‘Ž—‡ ‘ˆ ˆŽ—‹† ’”‡• ”‹„‡†Ǥ ‘™‹‰
–Šƒ––Š‡™‹‰Ž‡›Ǧ ‹ŽŽƒ’ƒ”ƒ†‹‰†‹†‘–‡š’Žƒ‹ Ž‹‹ ƒŽ‡š’‡”‹‡ ‡
‘”’—„Ž‹•Š‡††ƒ–ƒǡ ”‡ƒ†™‹†‡Ž›‘ˆŽ—‹†–Š‡”ƒ’›ǡ„—–‹–™ƒ•‘–—–‹Ž
ʹͲͳͲ –Šƒ–  ƒ‡ ƒ ”‘•• ƒ ”‡˜‹‡™ ƒ”–‹ Ž‡ ȏʹȐ –Šƒ– Ž‡† ‡ –‘ ”‡ƒŽ‹•‡ ƒ
™Š‘Ž‡ ‡™ ’ƒ”ƒ†‹‰ ˆ‘” ˆŽ—‹† ’”‡• ”‹„‡”• ™Š‘ ™ƒ–‡† –‘ „‡
’Š›•‹‘Ž‘‰‹ ƒŽŽ› ”ƒ–‹‘ƒŽǤ š’‡”‹‡–• Šƒ† ‘ˆ‹”‡† –Š‡ ”‘Ž‡ ‘ˆ –Š‡
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ˆ‘ —•‹ ”‡ƒ•‹‰Ž›‘Ž›’Šƒ–‹ ˆ— –‹‘Ǥdzȏ͵Ȑ

Š‡ British Journal of Anaesthesia ’—„Ž‹•Š‡† Š‡ ‡˜‹•‡† –ƒ”Ž‹‰

“—ƒ–‹‘ƒ† Ž› ‘ ƒŽ›š‘†‡Žƒ”ƒ†‹‰ˆ‘”ƒ–‹‘ƒŽ Ž—‹†”‡• ”‹„‹‰
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 Ž—‹†Š›•‹‘Ž‘‰› š˜

–Š‡”ƒ’›‹•— ŠŽ‡••‡ˆˆ‡ –‹˜‡–Šƒ™‹‰Ž‡›ƒ† ‹ŽŽƒ Žƒ‹‡†ǤŠ‡

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„‡‡–Š‡’Š‡‘‡‘‘ˆ‡š ‡•••‘†‹—•–‘”ƒ‰‡Ǥš ‡•••‘†‹—•–‘”ƒ‰‡
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™‹–Š‘—–‘˜‡”Ž›†‹Ž—–‹‰–Š‡‘š›‰‡Ǧ ƒ””›‹‰”‡† ‡ŽŽ•Ǥ ˆ™‡ Š‘‘•‡–Š‡
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‡˜‹†‡–Ž› ”ƒ–‹‘ƒŽ ƒ† ‹ Ž—†‡† Š‡ƒ†ƒ Š‡ǡ ƒ—•‡ƒǡ ƒ† •‘‡–Š‹‰
ƒŽŽ‡††‡ ‘†‹–‹‘‹‰Ǥ

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ƒ”‡ ‡•–‹ƒ–‹‰ –Š‡ ’Žƒ•ƒ ˜‘Ž—‡Ǥ ‘‡ Šƒ˜‡ —•‡† ‘”ƒŽ ’Žƒ•ƒ
˜‘Ž—‡ƒ•–Š‡‹” Ž‹‹ ƒŽ‰‘ƒŽˆ‘””ƒ–‹‘ƒŽˆŽ—‹†–Š‡”ƒ’›ǡ‹’ƒ”–‹ —Žƒ”ˆ‘”
–”‡ƒ–‡– ™‹–Š ƒŽ„—‹ ‘” ƒ •›–Š‡–‹  ‘ŽŽ‘‹† •‘Ž—–‹‘Ǥ Š‡› –‡ƒ Š
ȋ™”‘‰Ž›Ȍ –Šƒ– ƒŽ„—‹ ‹• ‘•–”ƒ‹‡† ™‹–Š‹ –Š‡ ‹” —Žƒ–‹‰ „Ž‘‘†
˜‘Ž—‡ǡ‹‰‘”‹‰–Š‡ˆƒ ––Šƒ–‘•–‘ˆ–Š‡–‘–ƒŽ„‘†›ƒŽ„—‹‹•ƒ–ƒ›
‘‡ –‹‡ ™‹–Š‹ –Š‡ ‹–‡”•–‹–‹—Ǥ Š‡› –‡ƒ Š ȋƒ‰ƒ‹ ™”‘‰Ž›Ȍ –Šƒ–
ƒŽ„—‹ †‡–‡”‹‡• ’Žƒ•ƒ ˜‘Ž—‡Ǥ ”‘—† ͷΨ ‘ˆ ’Žƒ•ƒ ƒŽ„—‹
‘”ƒŽŽ›‡• ƒ’‡•–Š‡ ‹” —Žƒ–‹‘‡˜‡”›Š‘—”ƒ†‹•„ƒŽƒ ‡†„›ƒŽ„—‹
Another random document with
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industrious plants once more, with a tendency to stop having double
flowers.
There are one or two things about corollas that I am sure you
would like to know. One is, how did the flowers manage to change
stamens into corollas? Another is, how did they manage to give them
such bright colors?
About corolla-making,—if you are determined to know that, you
will have to take yourself off to that far-away time when there were
no flowers. Then, in course of time, while changing about and trying
to get fitted to their surroundings, the plants, as you know, rolled
some of their leaves into pistils and stamens. But still they had no
petals.
The pistils and stamens were flowers, however,—as much flowers
as they would ever be, no matter how much corolla they might
develop.
A corolla does not make a flower; by this time you know the
important part of a flower is the pistil and stamens, and so, even to-
day, some flowers, as the elms and some maples, have no petals at
all. When such maples are in bloom, you will see gay fringes
decorating the trees. This fringe is made of the long pedicels with the
stamens at the end. The stamens swing in the breeze, and the pollen
is blown to the stigmas which are often in flowers on different trees.
Now, as plants grew and adapted themselves to their
surroundings, they produced more seeds than could by any chance
find room in the earth to grow. So every little seed that fell had to
fight its way with a host of other seeds and plants. A defective seed or
a weak one would stand no chance at all. The others would crowd it
out. We know how that is in a garden. The delicate flowers have to be
helped or the strong weeds would kill them. We pull up the weeds
and let the flowers have the whole garden to themselves. But in the
woods and fields each plant has to take care of itself and struggle up
as best it can.
This fight of the plants for a place to grow in is called the struggle
for existence. Now, whatever would help a plant in the struggle for
existence would, of course, be of great benefit to that plant. As we
know, cross-fertilization is a very great help; it makes stronger and
 better seeds, and the plants whose
seeds were regularly cross-
fertilized would be the ones to
survive.
Where pistils and stamens are
forming, there is a great deal of
nourishment brought to that part
of the plant, and substances are
being changed there. Very often
sweet juices are present. Long ago
when insects, in flying about,
smelled these sweets they
doubtless would go and eat them,
and they would also eat the pollen.
As they went from flower to flower
looking for food, they would carry
pollen sticking to their legs or
bodies, and so would sometimes
fertilize the flowers.
The seeds from such flowers
would be strong and would have
the best chance to survive. The
plants that grew from these seeds
would also inherit the tendency to
secrete sweet juices near the
flower.
In probing for sweets, the insect
would irritate the parts it touched,
and this would cause an extra flow
of sap there and very likely the
manufacture of more sweet juice; so the nectary came to be
developed.
You can understand how this might be by recalling how the skin of
your hand changes when you first try to do some new and hard work,
like rowing a boat.
After you have rowed a little while your hand is blistered. The
constant rubbing of the oar in one place has irritated it, just as you
can imagine the tongues of the insects rubbing against the delicate
flower tissue would irritate it. Wherever a place on the skin is
irritated, the blood flows to that spot; and so in the plant, where it is
irritated, there will likely be a collection of sap. After the blood has
flowed to the place on your hand which was rubbed by the oar, the
spot becomes red and inflamed and pains you, and finally the skin
separates in the form of a blister and a new skin forms underneath;
and if you keep on rowing, your hand does not keep on blistering, but
actually makes a new kind of skin to protect the rubbed places, and
what we call a “callous” or hard spot is formed. The skin is many
times thicker here than elsewhere, and was formed on purpose to
protect the place. So we can understand how irritation might change
a plant organ and in time form a nectary.
But how about petals, you are asking. Well, imagine yourself in
those old times when plants made their first flowers out of pistils and
stamens only.
These primitive flowers were probably not very showy. Primitive
flowers means first flowers,—flowers that lived way back in the
beginning of plant life.
They had no petals, but they secreted juices which the insects
liked. Those early insects were queer fellows, too, not very much like
our insects, except that they were fond of sweets and liked to eat the
tender parts of the flowers, just as our insects do to-day. They ate
nectar when they could find it and did not disdain pollen, which, it is
to be feared, they sometimes ate, anther and all; and, what is worse,
they in all probability frequently dined on pistil, which was very bad
for the plant.
Now imagine one strong plant secreting a good deal of nectar. The
insects would be likely to eat this and let the pollen and pistil alone,
only in getting to the nectar, they would be apt to dust the pistil with
pollen from another plant which they had been visiting and would
also brush off some pollen against their bodies.
Thus the strong plant with the abundant nectar would be cross-
fertilized and would keep its pistil unharmed. It would be very likely
to develop good strong seeds that would grow and again bear strong
flowers with plenty of nectar. Now, remember the essential organs—
that is, stamens and pistil—seem to find it a little easier to change
than other parts of the plant; so it would not be surprising if in time
some of the stamens were to become different. You see, the insects in
visiting the flowers would irritate them more or less walking over
them and clinging to them, and they would be likely to undergo
change for this reason; and if it happened that in some flower a row
of stamens got too full of sap to know what to do with themselves
and so spread out a little broader and more leaf-like and kept their
yellow stamen color or bleached-out white, that flower would be seen
far and near and the insects would go straight to it, for insects have
the sharpest kind of eyes for seeing bright colors a long way off. You
see what would happen; all the flowers whose stamens had done so
would be abundantly cross-fertilized,—that is, all their seeds would
get fresh pollen from another strong plant, and the plants growing
from these seeds would inherit the tendency of their parents to form
petal-like parts from some of the stamens. The flower could well
afford to lose part of its stamens for this purpose. Of course as time
went on, these stamens, which were half petals, might develop more
and more in the direction of signals,—that is, might become more
and more perfect petals, finally losing all trace of their old life as
stamens.
Of course no one can say that is just the way it came about, but it is
likely that in some such way it happened, for there are proofs of it
which you may like to read when you grow older.
So, you see, flowers are nothing but leaves after all,—very much
changed leaves, to be sure, but yet just leaves.
Sometimes when plants and animals have changed into a new
form, they change back again. We know some plants which once had
petals but which have again lost their petals and gone back to a form
which has no petals. Such backward changes we call retrogression,
and it is sometimes difficult to find out whether a flower with no
petals is a primitive form which for some reason has not changed or
whether it is one which has changed and gone back again. Usually,
though, we can find traces of petals and sepals in flowers which have
retrogressed.
You see, a flower depends upon its surroundings for its shape. If
its surroundings (and of course this includes its insect visitors) are
such as to favor its growth in the line of petals, it does so. But if for
some reason it becomes easier for it to grow and be fertilized in some
other way, perhaps by making abundance of light pollen which is
blown by the wind, as in the maple trees, then it may gradually lose
its petals, as it depends less and less on insects and more and more
on the wind for cross-fertilization. Nothing in life stands still; it is
always moving,—going on or going back. And this, we know, is just
the same in human life.
 We cannot stand
still; we must keep
growing wiser and
stronger and better,
or else we must do the
opposite.
 SIGNS OF OTHER TIMES.
In the beginning flowers seem to have had
their petals all separate from each other. Some
do still, and these we call polypetalous,
because “poly-” means many, and they have
many petals. But other flowers, like our
morning-glory, have no separate petals; all are
grown together into a tube with a bright
border.
But this tube and border tell us a little story
if we are able to hear it.
They tell us of the time when the morning-
glory had several petals. More than this, they
tell us just how many it had. If we were to
guess we should probably say five, because it
seems so fond of the number five, with its five
nectaries, five nectar guides, five stamens, and
five sepals.
If we guessed five we should guess just
right. There is no doubt but that once upon a
time the plants from which our morning-
glories are descended had five separate petals.
The morning-glories themselves manage it
differently now, but it took them a long time
to do it. They were working away, long before
the great pyramids of Egypt were built, to get
their five petals united into one piece. But it is
done, and they have learned how to twist the
flower up tightly in the bud and then unroll it
in all its glory.
They never have five petals now, but they still bear traces of it.
Look at the little notch on the border, halfway between two nectar
guides. Does that tell us anything?
Count the notches. Five, you see.
 Look at the line that runs from the notch down
to the bottom of the flower.
The corolla looks as though it had been folded
along those lines. You can easily see five long
creases ending in a notch. The flower is folded
along these lines in the bud, but we think the lines
have yet another meaning.
Carefully tear the corolla down the lines; you
see, a very little pressure does it. Now we have the
corolla in five parts, like five petals, only it is so
weak it can no longer hold itself up. Once upon a
time we think it grew this way, with five separate
petals, only the petals stood up then, for they must
have been stiffer and perhaps were not so long. It was long, long ago,
oh, very long ago, that it had its five petals. Then the edges of the
petals began to grow together, and they kept on doing this until, in
course of time, the whole length of each petal had grown fast to the
next one, all except that little tiny spot where the notch is.
We are glad our morning-glory kept this little notch and the line
where the sides of the petals grew together, for that is what tells us
the story of long, long ago when all the petals were separate.
When finally they were grown together, the corolla did not need to
be so stiff, for its shape helped to make it firm, and then it no longer
used good material to make stiffening for the petals, for that would
have been a waste of plant sap, and plants do not like to waste
materials. When they find they can get along without something they
have been used to having, they stop making it. Life is too short and
too precious to waste a bit of it. Our flower only kept the stiffening in
the corolla along the paths where it wished the bees to go to its honey
cups and where, when folded, it could best protect the bud.
The morning-glory, you see, is as wise as it is beautiful.
 WHY ARE THE FLOWERS SO LARGE AND
BRIGHT?
Why are the flowers so large and bright?
We cannot say that they
were always so. It is probable
they were not. But good
Mother Nature has watched
over them as they came upon
the earth, and she has
lovingly made them so large
and bright.
How could she do this? Let
us see. Here is a tangle of
plants. They all bear flowers
and all set seeds. Some are
stronger and more beautiful
than others. The seeds fall to
the ground. Those from
strong and beautiful plants
are larger and stronger than
the others. After a while the
seeds sprout. Not all do this,
however. The very weakest
do not sprout. Dear Mother
Nature has other work for
them. “You are not suited to
struggle in the earth with the strong seeds, dears,” she whispers and
lays them to rest. They do not wake up; the materials in them
change. These materials let go of each other; they depart from the
seed; some as gases float off in the air; others as minerals sink in the
earth. The gases and the minerals are not lost. They join some other
plant and help to make it strong.
“It is better to help another than to try to grow yourselves,” Mother
Nature whispers to these little seeds that could not sprout. And they
are happy. They are glad to change into gases and minerals and help
another plant to grow.
Many of the seeds sprout, but not all grow up and blossom. There
is not room in the earth for all the seeds to grow; there is not food
enough in the air to feed so many. Mother Nature with her kind eye
looks over the growing plants.
She smiles and shakes her head at those trying to grow in shady
places.
“No, dears,” she whispers, “there is other work for you to do.” Then
the shaded seedlings do not try any more to grow into plants. They
give up the materials they have collected to the little brothers and
sisters who have started in the good ground and the sunlight.
They fade away, but they are happy, for they, too, are doing their
work. The materials in them let go of each other. They change into
gases and float off in the air, or to minerals and other substances and
sink to the ground. These gases and solid substances pass into other
plants and help make them strong.
“It is better to help another than to do poor work alone, dears,”
Mother Nature whispers, as she lays them to rest.
Then she visits all the weak plants, and all those in poor soil or in
too much light or too much shade, and lays them to rest. Their
materials go to nourish the strong plants, who are doing good work
in the world and growing in beauty. Not all the plants that live to
blossom are good alike. Some are better than others, but Mother
Nature lets them grow if they are strong enough and can find food.
At last the blooming comes. The flowers do their best. The strong
ones make large, bright flowers full of color and full of sweetness.
Mother Nature smiles at them and is pleased. The weaker flowers do
their best; they are not so bright nor so large. Mother Nature smiles
at them, for she loves them, too, and she will tell them what to do.
The bees come and fly to the brighter flowers; they have rich,
abundant pollen and rich nectar. The bees know this; they do not
care so much for the duller, smaller flowers.
When the bees do not come, Mother Nature whispers to the little
flowers, “Never mind, dears, there is work for you to do.” So they are
happy, though their ovules get no pollen and they set no seeds. They
are happy to do the work dear Mother Nature has for them to do.
 The strong
flowers set
their seeds;
they are
strong, and
they have
been well
fertilized.
The weak
flowers set
few seeds;
they are not
strong to
make many
seeds, and
they have
not been well
fertilized. So
year by year
and century
by century
Mother
Nature
watches her
plants and
encourages
the strong to
grow and helps the weak to find other work.
And this is why the flowers are so bright.
Mother Nature selects those that are to grow and blossom and
sends the rest to help them. This is what we call natural selection,
and this is what makes the earth so beautiful. Only the best continue
to grow; the others are glad to help them.
HOW MOTHER NATURE MAKES NEW FLOWERS.
Once upon a time there lived a little plant in a marshy place. We
will call it Primus, not because that was the very first form of the
plant, for it was not, but because that was its form when we first saw
it.
It had five small yellow petals, five small stamens, and an ovary.
When its seeds were ripe, along came a great wind and blew them
away from the marsh upon the dry land at the edge.
Poor little seeds, they were out of their familiar wet marsh and
they could not grow. But they did their best. Some of them managed
to sprout, but soon they found the earth too dry and the sun too hot;
so they said, “We will turn to other work; we will help the other
plants and not try to grow ourselves.”
So they changed into gases and minerals and other substances. But
a few of the seeds continued to grow.
They blossomed and bore seeds, but they were not just like the
plants in the marsh. Mother Nature had helped them get a tougher
skin and taught them how to shut tightly their pores in dry weather,
so that the water within them could not escape.
You see, they were already different from their parents, though you
might not have noticed it if you had seen them, the difference was so
slight. The seeds of these new plants sprouted the next season. They
did not have a hard time to grow. They knew just what to do, and the
best and strongest of them grew a few hairs to help cover up the
pores, so the water would not go out too fast.
It happened to be a very hot, dry season, and all the plants but
these hairy ones stopped growing. They changed into gases and
minerals and other substances to help the other plants. The hairy
people got through the dry season very well. They set a good many
seeds, and these seeds sprouted. The new plants remembered about
the hairs and had plenty of them. Some were covered all over with a
soft down.
 And it was well they were, for it was a very hot, dry season, and all
but the downy ones stopped growing and changed into minerals and
gases and other substances to help the others. The seeds of the
downy plants blew far over the dry land, far away from the marsh;
but they had learned to live in the dry soil, and if you had found these
downy people, you would hardly have known they were descended
from the smooth, juicy, large-leaved marsh plants. Their stems were
hard and tough and their leaves stiff and small. We can no longer call
them Primus, they are so changed.
Let us call them Secundus. Secundus had small yellow flowers, like
the marsh plants it was descended from. But one day some of the
seeds of Secundus blew into the edge of a wood where the soil was
rich and the air damp. This just suited the Secundus seeds, and they
grew into very thrifty plants indeed. They had so much sap and grew
so luxuriantly that their petals were twice as large as was usual with
Secundus petals. These fine showy flowers also possessed a great
deal of nectar, they had so much sap. Of course the bees came to
them, and they were well fertilized. They set many seeds. The next
year these strong seeds were able to grow even when their neighbors
were not, and the plants that came from these seeds also had large
showy flowers.
These stronger plants held their own, you may be sure, and at last
there was more of them than of the small-flowered plants. It was well
for them this was so, for there came several bad seasons when
nothing was just right for these plants. It was cold and stormy, and
only the very strongest lived through it. But they managed to survive,
and their flowers were large and showy.
All the weaker plants with smaller flowers were killed out, and only
these large-flowered ones remained. They were very different from
their ancestors the marsh plants, and we shall have to call them
Tertius.
One day some of the seeds of Tertius were blown into a new kind
of soil; they sucked up the juices of this new soil, and lo! some of
their flowers opened white instead of yellow. It so happened that the
white-flowered plants were stronger than the others. The bees liked
them, too; for, being so strong and full of sap, they made plenty of
honey. So these white-flowered ones increased in numbers very
greatly. At last only the white ones could be found; the yellow ones
had gradually given way before them until no yellow ones were left.
So we will call the white-flowered people Quartus.
Quartus lived a long time, each year bearing seeds, the strongest
and best of which grew up and bore flowers.
One day some of Quartus’ seeds were blown into a hot, sandy
place; this almost killed them, but some of them managed to grow.
Their leaves were smaller and stiffer than ever before, but they had
a great many of them, and their flowers were large and white. They
grew to like the sandy soil, and what they got from it changed their
sap in some way so their petals were delicately tinged with pink. The
bees liked these pink flowers; perhaps their honey was a little richer;
perhaps they could see them better. However that may be, the bees
almost deserted the white-blossomed plants and visited the pink
ones. So the white flowers set few seeds and the pink flowers many.
When the seeds sprouted, the pink ones were the strongest, because
in their change of color there was somehow added a change in
strength; they were stronger than the white flowers. They grew fast
and took the materials from the earth and the air; and when the
white flowers saw this, they said, “It is their turn now,” so they
changed into gases and minerals and other things and helped the
pink flowers to grow.
Soon there were no more white flowers to be seen; they had
stopped growing, and only the pink ones kept on, so we shall have to
call these pink flowers Quintus.
But a great danger threatened Quintus. Cows and goats and sheep
bit off their leaves. They ate so much of them that many plants were
killed outright. Only the stiffest and hardest were left to blossom and
set seed. The seeds of these plants with the stiff leaves and stems
grew into other stiff-stemmed and stiff-leaved plants. The cattle
browsed the tenderest of these and again left the stiffest. This went
on for many years, the plants growing stiffer and harder each year.
Some of them got so stiff and hard that they threw out prickles all
over their stems.
These prickly ones were not eaten, and in time you would have
found them grown into woody bushes with prickly stems.
We shall have to call these Sextus.
 Sextus spread all over the sandy plains. Hardly any other plant was
to be seen. The strong Sextus seeds sprouted and took the materials
in the earth and the air, and the other seeds that happened to be
blown among them did not grow; they changed into gases and
minerals and other substances and helped the Sextus plants to grow.
One day some Sextus seeds blew upon good, rich, damp soil, and
there they sprouted and grew. They had plenty of water, and there
were no cattle to disturb them; so those with the fewest prickles were
the best off, because they could use the food material to make larger
flowers instead of prickles. So the plants with fewer prickles had
larger flowers and better seeds, and these seeds sprouted and grew,
and the others gave way before them. In the course of time these
plants growing on the rich soil lost their prickles, and their flowers
were large and very deep pink; in fact, some of them were a bright
red.
These bright red flowers attracted the bees, and so they lived on
and set seed. These we must call Septimus.
For some reason some of the seeds of the Septimus flowers
developed unusually thrifty plants.
These plants had flowers with petals so full of sap they overlapped,
and finally, just because they were so full of the growing spirit, the
edges of the petals grew together.
Finally, the flowers with the edges grown together were the most
successful. The tube their flowers made kept the nectar for the bees,
and the bees liked to go into these red bells. You see what had
happened: the flowers were no longer polypetalous. Their petals had
grown together; they were gamopetalous. Their corollas formed snug
tubes, something like a morning-glory corolla, for the bees.
We shall have to call these people Octamus.
And we will not follow them any farther, only be sure they kept on
changing ever and ever. Whenever the seeds fell in a new soil, they
had to change or die. The reason they could change so is because no
two things are ever just alike, and out of a great many plants some
might be fitted to survive in the new surroundings. These would live,
and their descendants would be like them, but they would be
different from their ancestors.
 In some such way, no doubt, the many different kinds of flowers
have come into existence.
If you ask me for the exact name of our plant that has changed so
many times, I cannot tell you, for I do not know.
But that, we believe, is Mother Nature’s way of making new
flowers.
 TONGUES AND TUBES.
A flower tube is a most convenient and
safe place to keep stamens and nectar. If
it is protected by scales or hairs or a
sticky juice, as is often the case, the ants
and other small insects are given a
gentle but convincing hint to keep out.
They might readily infer their presence
is not wanted, and though it may hurt
their feelings a little, they have nothing
to do but obey.
Some flowers like ants and little
crawling insects, but they have open,
spreading corollas with the nectars
easily reached; but you may be sure a
flower with a tube is no friend to them.
Its tube says “keep out” as plainly as
though it had put out a printed sign, and
then a tube is a sign anybody in the
insect world can read, no matter what
language he may speak or whether he
knows his letters.
But tubes are not intended to keep all
visitors away,—far from it.
They are as much an invitation to one kind of insect as they are a
request to “keep off these premises” to another. If you happen to be a
large insect with a long tongue, you will be sure to find a welcome in
many a flower with a tube. And no doubt, if you are fond of honey
and are industrious about collecting it, you will find that the flower
whose nectar you like the very best and which you visit the oftenest
has a tube just the same shape and size as your tongue; and what is
more, it will be in the most convenient position for you to reach it.
It seems to be your flower, and no doubt it is, for flowers have a
way of making their tubes to fit the tongues of those who love them
best. Not that they do all the fitting, for no doubt the tongues also
grow to fit the flowers.
Of course other insects with similar tongues can get the honey too,
and a good many, whose tongues are quite different, can reach more
or less of it; but the bulk of the honey is for the favorite visitor. He
can reach clear to the bottom of the nectary, and in some cases,
where the favorite insect has a very long and very slender tongue, the
spur, or tube, will be so long and slender that none but that
particular kind of insect can get the honey at all.
Everybody who lives in New England, and a good many who do
not, knows the white azalea, often called swamp honeysuckle.
Swamp honeysuckle and the large night-flying moths are great
friends. The azalea has provided honey for the fellows, and protects
it, too, against other visitors, all but the bees and humming birds.
The humming birds are welcome, and the bees have a way of coming
whether they are welcome or not.
If you go just at dark to where the azaleas are blooming, you will
not see the moths, but you will hear them. The chief sounds in the
woods are the rustling of twigs and leaves in the breeze, the calling of
frogs from the ponds, the noises of the insects, and the voices of the
night-flying birds. Then all at once there comes another sound,—a
steady buzz-z-z that draws nearer and nearer until it seems to be
close to your ear. This is the moth come to visit the honeysuckle.
And, no doubt, the honeysuckle is glad to feel the breeze of these
fanning wings and feel the long tongue enter the tube, for the moth’s
body touches the out-reaching stigma and leaves there pollen from
some other flower whose honey it has enjoyed. From the stamens it
detaches pollen grains to carry to another flower; and this, too, no
doubt, gives happiness to the azalea, for it makes its pollen, not for
its own use, but for the sake of its azalea friends.
You see, the azalea has long, upturned filaments that reach far out
of the tube, and the style is yet longer, so that only a large insect or a
humming bird, collecting honey while on the wing, can really give
pollen to the stigma.
Bees alight back of the anthers and take the honey. If they want
pollen they collect it from the stamens without touching the stigma,
except once in a while by accident, as it were. So however much the
majority of flowers may love and respect the bee, our azalea has no
liking for her. Besides, the bee has a bad habit of biting a hole in the
flower tube and getting the honey that way. This would be a
thoroughly disreputable performance on the part of any insect, and if
bees are not ashamed of it they ought to be.
The azalea does several things for the moth it loves. It may be its
beautiful white color is for his sake; anyway, if the flower were not
white the moth would not be likely to find it, since he flies abroad
after the birds have gone to rest,—that is, in the evening, when it is
dark in the damp thickets where the honeysuckle loves to grow.
Azalea has a sweet white corolla with a long, slender tube containing
nectar that moth or humming bird can reach, but which bees cannot
reach. Watch a bee try some time. If the flower is between you and
the light, you can see the bee’s brown tongue through the flower
tube; she appears to be standing on her toes and reaching in as far as
she can; she darts out her tongue to its full length, and you can see it
wriggling and straining to get to the abundant honey low down in the
flower tube. But there is no use trying; the tongue is too short and
the tube too long. The honeysuckle tube was not made to fit the bee’s
tongue, and the bee can get only the outer rim of the honey. Perhaps
this is why the bee so often breaks in the back way.
Besides being white, the azalea flowers grow in clusters, which
makes them yet more visible in the dusk. They exhale a delicious and
far-reaching perfume too, and this is a note of invitation to the
moths.
Instead of writing a note on a
sheet of perfumed paper, the
honeysuckle simply sends the
perfume without the paper, and the
moth understands the message and
knows the white azalea “requests the
pleasure” of his company that
evening, and he puts on his best
manners, since he cannot change his
clothes, and goes.
The white azalea is so very sweet
and so pretty, it would not be
strange if other uninvited guests
than bees were to visit it. No doubt,
the ants and bugs and gnats and flies
would be glad to, but the azalea has
a very inhospitable way of receiving
such would-be guests. All over the
outside of the lower part of the white
tube and running in a line to the
very tips of the petals are tiny white hairs with black tips.
These are azalea’s body guard. Each tip exudes a drop of sticky
liquid.
Fine, sticky hairs cover the stems and the leaves too; so the
unfortunate insect that tries to crawl up to the flower is sure to get
wings and legs hopelessly entangled and stuck together.
Only large fellows, like bees, who are strong enough to pull
themselves free and clean off their legs, are able to defy this body
guard. You will sometimes meet our sweet azalea covered on the
outside with little marauders who wanted to steal her honey but
could not, because the body guard caught them and stuck them fast.
Not all flowers with tubes have succeeded as well as azalea in
keeping their honey for the visitors who can do them the most good.
Yet many have tried.
Look at the morning-glory, for instance; it has hairs at the
entrance to the nectaries which the ants cannot readily pass, but
which the bees can push aside. The openings to the nectary are large
enough readily to admit the tongue of a bee, and the distance into the
nectar is about the length of a bee’s tongue; but there are no sticky
guards to preserve the honey, for the bees and small beetles and
other tiny insects often crawl into the tube and eat the honey and
even devour the flower itself.
 Evening Primrose.

Tropæolum has a fine large tube full of rich honey for bees and
humming birds. This tube no doubt corresponds to some tongue or
bird-bill in her own South America. But in our country the bees
answer very well. The bumblebee is fond of Tropæolum honey and
fertilizes the flower, while an occasional ruby throat may be seen
taking a sip.
Jewelweed’s horn is a humming bird tube and a bee tube, too. The
flowers are so delicately balanced on tiny stalks that wingless insects
would not find an easy entrance.
Pelargonium, too, has a tube suited to some long and slim-tongued
visitor. In her own native land in far-away Africa she probably loves
the butterflies that live there, who also love her, and so they have
grown tongue and tube to fit each other. For the flower is not the
only one to change: the insect changes to suit the flower at the same
time that the flower changes to suit the insect. They grow to fit each
other.
Wherever you see a flower tube you may be sure there is
somewhere a tongue to fit it.
 GLOSSARY.

L. = Latin. A.-S. = Anglo-Saxon.

 A.
Acheloüs, n. A river god with whom Hercules wrestled. Like
Proteus, Acheloüs could change his shape; he became a serpent
and a bull, but Hercules vanquished him nevertheless and tore
off his horn, which became the horn of plenty.
Alternate, a. L. alter, another; one following another. Said of
leaves standing singly at the nodes of a stem; also of stamens that
stand between the petals, and of petals that are placed between
the sepals.
Amalthea, n. In Greek mythology, the nurse of Jupiter, probably
a goat.
Amœba, n. From a Greek word meaning “change”; the name of
one of the lowest forms of life; a bit of living protoplasm capable
of existing as a single cell and of changing its form at will.
Ancestors, n. L. antecessor, a foregoer; forefathers; those from
whom animals or plants are descended.
Animal cells, n. The cells or minute divisions which make up the
animal body.
Animals, n. All living things which are not plants are animals. In
the lower forms of life it is impossible to decide whether certain
living things are animals or plants.
Anther, n. From a Greek word meaning “flower”; that part of the
stamen containing the pollen.
Anther cells, n. The hollow spaces in the anther where the pollen
is kept.
Aristocrat, n. From two Greek words meaning “best” and “rule”;
one belonging to the best in a community; one among those fit to
rule.
Aristocratic, a. Like an aristocrat.
Axil, n. L. axilla, little armpit; the angle formed between the upper
side of a leaf and the stem or branch to which it is attached.
Azalea, n. The name of a plant. The “swamp honeysuckle” is not a
honeysuckle, but is an azalea.
 B.
Barb, n. L. barba, a beard; a tuft of hairs; a sharp point projecting
backward from the point of a fish hook or arrow or any other
sharp-pointed instrument. The barb prevents the instrument
from being readily withdrawn.
Bark, n. The outer covering of the stems and roots of woody
plants.
Beak, n. The bill of a bird; the long, projecting point in the fruit of
the geraniums.
Bloodroot, n. An early spring flower. A pretty, delicate, white
flower opens on a stem that comes up from the ground, and the
roots, when wounded, yield a blood-red sap.
Boer, n. D. boer, a farmer; a peasant; the name of the Dutch
colonists of South Africa. They are principally farmers and cattle
raisers. They have had many difficulties with the English settlers,
in some of which blood has been shed.
Bract, n. L. bractea, a thin plate of metal; gold-leaf. Used of small,
usually thin, leaf-like parts, and often found near a flower or
flower cluster.
Bulb, n. L. bulbus, a bulbous root; an onion; the name of the
underground, scale-covered part of hyacinths, etc.
 C.
Cactus, n. From a Greek word meaning “a prickly plant”; a group
of plants which usually grow in dry places and have prickles or
thorns instead of leaves. The prickly pear grows wild in northern
latitudes, and others, such as the night-blooming cereus, are
often seen in hothouses.
Callous, a. L. callosus, hard-skinned, thickened and hardened.
Applied to a hard place on the skin, usually the result of friction.
Calyx, n. From a Greek word meaning “to cover”; the outer set of
envelopes which form the perianth of a flower. If the perianth
has but one set of envelopes it is called the calyx.
Capillary attraction, n. The force which causes liquids to
disperse through fabrics or tissues. If one end of a towel be
placed in a bowl of water, the whole towel will be wet in course of
time.
Carbon, n. L. carbo, a coal; a substance very widely distributed
and existing under various forms. Coal is one form of carbon,
graphite another, the diamond a third. One atom of carbon
combined with two of oxygen form carbon dioxide.
Carbon dioxide, n. A heavy gas, found as an impurity in the air.
It is breathed out by animals and plants, and absorbed and used
as a food by plants.
Castor oil, n. The oil obtained from the seeds of the castor-oil
plant. Used as a medicine and also in dyeing cotton certain
colors.
Cell, n. L. cella, a small room; a case or cup in which something is
held, as anther cell, ovary cell, honeycomb cell; also the
protoplasmic particles of which plants and animals are built up.
Candelabrum, n. L. candela, a candle; a candle stick; any
branched candlestick. A candelabrum rests on a post, while a
chandelier is suspended. Candelabra is the plural.
Chasm, n. From a Greek word meaning “a yawning hollow”; a
wide, deep cleft.
Chlorophyll, n. From two Greek words meaning “light green” and
“leaf,” leaf-green; the green coloring matter of vegetation.
Columbine, n. L. columba, a dove; a flowering plant which gets
its name from the fancied resemblance of its petals and sepals to
the heads of doves round a dish.
Complexity, n. L. com, together, plectere, to weave; formed by a
combination of simple things.
Convolvulaceæ, n. The name of a family of plants to which
belong the morning-glory and bindweed.
Cornucopia, n. L. cornu, horn, copia, plenty; horn of plenty.
Corolla, n. L. corolla, a little crown; a garland; the floral envelope
within the calyx, very often bright colored.
Cotton-seed oil, n. An oil expressed from the seeds of the cotton
plant and, when purified, used instead of olive oil.
Crete, n. An island to the south of Greece.
Crocus, n. An early spring flower.
Cross-fertilization, n. The fertilization of the ovules of one
flower by the pollen of another.
Cross-fertilized, a. Fertilized by the pollen from another plant.
 D.
Dew, n. The moisture of the air when condensed on any cold
surface. Dew does not fall; it is formed wherever moisture in the
air comes in contact with a substance colder than the air. Hence
there may be dew on the under as well as the upper side of a leaf.
Dissolve, v. L. dis, apart, solvere, loose; to separate the solid
particles of a body in a liquid; to melt. Sugar dissolves in water.
Double flowers, n. All those whose petals are numerous.
Sometimes the stamens are changed into petals, as in double
roses, and sometimes even the pistils have become petals.