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University of Groningen North Sea seaweeds: DIP and DIN uptake kinetics and management strategies Lubsch, Alexander

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University of Groningen

North Sea seaweeds: DIP and DIN uptake kinetics and management strategies Lubsch, Alexander

IMPORTANT NOTE: You are advised to consult the publisher's version (publisher's PDF) if you wish to cite from it. Please check the document version below.

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Publication date: 2019

Link to publication in University of Groningen/UMCG research database

Citation for published version (APA):

Lubsch, A. (2019). North Sea seaweeds: DIP and DIN uptake kinetics and management strategies. University of Groningen.

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North Sea seaweeds:

DIP and DIN uptake kinetics and management strategies

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The research reported in this thesis was carried out at the Department of Estuarine and Delta Systems (EDS) at the Royal Netherlands Institute for Sea Research (NIOZ), Texel, supported by the

Faculty of Science and Engineering at the University of Groningen.

Lay-out: Alexander Lubsch Cover design: Alexander Lubsch

Printed by: MünsterMedienXpress GmbH & Co. KG ISBN: 978-94-034-1688-5

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North Sea seaweeds: DIP and DIN

uptake kinetics and management

strategies

Phd thesis

to obtain the degree of PhD at the University of Groningen

on the authority of the Rector Magnificus prof. E. Sterken

and in accordance with

the decision by the College of Deans. This thesis will be defended in public on

Friday 24 May 2019 at 14.30 hours

by

Alexander Lubsch

born on 14 October 1980 in Lengerich, Germany

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Supervisors

Prof. K.R. Timmermans Prof. T.J. Bouma

Assessment Committee

Prof. K. Bischof Prof. A.G.J. Buma

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i

Preface

The term ‘seaweeds’ is used here to describe multi-cellular marine macro-algae, although at some stage in their life-cycle seaweeds can be unicellular (spores, zygotes). One key component for survival and growth in seaweeds is the availability of (dissolved inorganic) nutrients. In this study the term ‘nutrients’ refers to the biologically available inorganic dissolved forms of nitrogen and phosphorus, two major constituents of living organisms. The focus in this thesis is on investigation of nutrient uptake kinetics, nutrient management, effects of nutrient limitation, growth, and composition in relation to nutrient availability in 4 ecologically and economically relevant (green, brown, and red) native North Sea seaweed species: Ulva lactuca (Chlorophyta),

Saccharina latissima, Laminaria digitata (Phaeophyta) and Palmaria palmata (Rhodophyta). Insight in

these major stimuli of production in seaweed biomass will allow better understanding of the ecophysiology of seaweeds, both under natural, as well as (large scale) production conditions.

It can be envisioned that nutrient availability will not only determine growth, but through the effects on composition it may also affect colour and texture. Hence, in addition to the analysis of the nutrient uptake kinetics and nutrient management strategies, 2 novel approaches with ecological and economical relevance on seaweeds with regard to nutrient availability are presented. Firstly, a combination of spectrophotometric measurements and colorimetric techniques was applied to determine the colour appearance of U. lactuca, resulting in a smartphone application ‘EyeOnUlva’. This application evaluates the nutritional value (total dissolvable protein content) of this opportunistic seaweed by digital imaging. Secondly, an approach was introduced that allows for standardised methods to infer to effects of nutrient availability and varying hydrodynamic forces on the texture of seaweed, using L. digitata individuals.

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ii

Contents

Chapter 1 ... 1

1. General introduction ... 1

1.1 Seaweed... 1

1.2 The North Sea ... 2

1.3 Seaweed farming in Europe ... 5

1.4. Thesis outline ... 8

Chapter 2 ... 26

Uptake kinetics and storage capacity of dissolved inorganic phosphorus and corresponding N:P dynamics in Ulva lactuca (Chlorophyta) ... 26

2.1 Abstract ... 26

2.2 Introduction ... 27

2.3 Material and methods ... 29

2.4 Results ... 35

2.5 Discussion ... 41

2.6 Acknowledgements ... 46

Chapter 3 ... 47

Using a smartphone app for the estimation of total dissolvable protein concentration in Ulva lactuca Linnaeus (Chlorophyceae) ... 47

3.1 Abstract ... 47

3.2 Introduction ... 48

3.3 Material and methods ... 51

3.4 Results ... 57

3.5 Discussion ... 63

3.6 Acknowledgements ... 67

Chapter 4 ... 68

Uptake kinetics and storage capacity of dissolved inorganic phosphorus and corresponding dissolved inorganic nitrate uptake in Saccharina latissima and Laminaria digitata (Phaeophyceae) 68 4.1 Abstract ... 69

4.2 Introduction ... 69

4.3 Material and Methods ... 73

4.4 Results ... 80

4.5 Discussion ... 90

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iii

Chapter 5 ... 98

Texture analysis of Laminaria digitata (Phaeophyceae) thallus reveals toughness gradient along lamina ... 98

5.1 Abstract ... 98

5.2 Introduction ... 99

5.3 Material and methods ... 101

5.4 Results ... 107

5.5 Discussion ... 110

5.6 Acknowledgements ... 115

Chapter 6 ... 116

Dissolved inorganic phosphate uptake and corresponding dissolved inorganic nitrate uptake in the seaweed Palmaria palmata (Rhodophyceae): ecological and physiological aspects of nutrient availability. ... 116

6.1 Abstract ... 116

6.2 Introduction ... 117

6.3 Material and methods ... 120

6.4 Results ... 125

6.5 Discussion ... 134

6.6 Acknowledgements ... 141

Chapter 7 ... 142

‘Manual for nutrient uptake kinetics in seaweed cultivation’ ... 142

7.1 Offshore cultivation ... 142

7.2 Integrated multi-trophic aquaculture (IMTA) ... 145

7.3 Tank cultivation ... 148

7.4 Biofiltration ... 150

Chapter 8 ... 153

Synthesis ... 153

8.1 General findings ... 154

8.2 Innovative aspects / highlights of the thesis ... 159

8.3 Conclusions ... 160 8.4 Research outlook ... 161 References ... 164 Summary ... 191 Samenvatting ... 195 Acknowledgements ... 200

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